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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
O20 NDE,
1907, pp. 1-446.
(JANUARY—APRIL.)
PRINTED FOR THE SOCIETY —
20457438
AND SOLD AT THEIR HOUSE IN HANOVER-SQUARE,
LONDON:
MESSRS. LONGMANS, GREEN, AND €GQ,,
PATERNOSTER ROW.
Aces Sill
OF THE
COUNCIL AND OFF I@RREs:
OF THE
ZOOLOGICAL
SOCIETY OF LONDON.
WOOK
COUNCIL.
His Grace THe Duke oF BeprorD, K.G., President.
Grorce A. BouLencer, Esq.,
F.R.S., Vice-President.
JOHN Rose Braprorp, Esq.,
WILDS; IDESCh, IeHSh. oes
President.
F. Dawrrey Drewirr, Esq., |
M.A., M.D.
CHARLES Drummond,
Treasurer.
Str Epwarp Duranp, Br., C.B.
F. Du Cane Gopman, Ksq.,
D.C.L., F.R.S., Vice-Presi-
dent.
JOSEPH JACKSON ListER, Esq.,
M.A., F.R.S.
Sir EpmunpD Gites Lover, Br.,
Vice-President.
Pror. Epwarp ALFRED M1v-
cuin, M.A.
Ksq.,
|
P. CHatmers MitcHey, Esq.,
MA. (DIS coins SHS
Secretary.
W. R. Oaitvir-Grant, Esa.
ALBERT Pam, Esq.
E. Lort Puiwuirs, Esq.
Sir Parrick Puiayratr, C.1.E.
THE Hon. WALTER ROTHSCHILD,
D.Sc., M.P.
Howarp Saunpers, Hsq., Vice-
President.
Davin Seru-Smiru, Hsq.
OLDFIELD THomas, KEsq.,
A. Trevor-Battys, Esq., M.A.
Henry Woopwarp, Ksq., LL.D.,
F.R.S., Vice-President.
PRINCIPAL OFFICERS.
P. CHatmers Mircnrett, M.A., D.Sc., LL.D., F.BS.,
Secretary.
Frank E. Bepparp, M.A., F.R.S8., Prosector.
R. I. Pocock, F.L.8., Superintendent of the Gardens.
CHARLES
Pathologist.
F. H. Waternouss, Librarian.
JoHN Barrow, Accountant.
W. H. Cots, Chief Clerk.
GABRIEL SELIGMANN, M.R.CS.,
RAC as
GrorcGe ArtHur Dovusiepay, Clerk of Publications.
ARTHUR THOMSON, Assistant Superintendent of the Gardens.
LIST OF CONTENTS.
1907, pp. 1-446.
January 15, 1907.
The Secretary. Report on the Additions to the Society’s
Menagerie during the months of November and De-
Gemlboer WO OG spas acer eyeysclck a cacleee re rar ege ce pice eens ame sms
Mr. Oldfield Thomas, F.R.S., F.Z.S. Description of a new
Monkey from the Ituri Forest. (Plate I.) ...............
1. On a Collection of Mammals made by Dr. Vassal in
Annam. By J. Lewis Bonnore, M.A., F.LS., F.Z8.
(Bibi ire Ul) pears ARs SMR UB Oe ane ae aes enP One Aloe Sa Or Pose 03
2. On the “ Bleating” or ‘“ Drumming” of the ee
(Gallinago celestis). By P. H. Baur, B.A., F.ZS. .
3. Contributions to the Knowledge of the Systematic
Arrangement and Anatomy of certain Genera and
Species of Squamata. By Frank E. Bepparp, M.A.,
BUR: Se eerosecsorsortive: SoOcietige.s-6. dae ace eteen eee
4, A List of Moths of the Family Pyralide collected by
A. E. Pratt in British New Guinea in 1902-3, with
Descriptions of new Species. By Grorce H. Kenrick,
F.Z.S. (Plates IIT. & TRV SaaS ten tine Senin ten ce b ies oa ob
a 2
Page
12
30
68
1V
5. On some new and insufficiently known Species of Mar-
moset Monkeys from the Amazonian Region. By Prof.
Dr. Emit A. Gorupt, C.M.Z.S., Director of the Para
Museum
SO oe mee reese meee eee sees eseeeees es esaeeeesseseeresssersvsesee
February 5, 1907.
Mr. F. Martin Duncan. | Cinematograph exhibition of
Animals in the Society's Gardens and other Zoological
SIU OU I STCUS arlene ere tie) CRD CPP Ee AREA nA ShE let ac .509
Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition of a collec-
tion of Mammals and Birds from the Islands of Saghalien
and Hokkaido
Cem ee mew twee mes wesc eee neers ease eee se ese ese es teserene
Dr. W. T. Calman, F.Z.8. Notice of a paper on New or
Rare Crustacea of the Order Cumacea from the Collection
of the Copenhagen Museum
1; The Origin of the Lateral Horns of the Giraffe in Fetal
Life on the Area of the Parietal Bones. By E. Ray
Lanxester, M.A., D.Sc., LL.D., F.R.S., F.Z.S., Director
of the Natural History Departments of the British
Museum
2. Parallel Hair-fringes and Colour-striping on the Face of
Fetal and Adult Giraffes. By E. Ray Lanxsster,
MA’, DSc., UE.D., EF.R.S:, F:Z.8., Director of the
British Museum (Natural History). (Plate V.)
3. On the Existence of Rudimentary Antlers in the Okapi.
By E. Ray Lanxester, M.A., D.Sc., LL.D., F.R.S., F.Z.S.,
Director of the British Museum (Natural History).
((Bbites VAI GG BY Wi th PP Ok He kine ah a aA
4. Description of Hyla resinifictrix Goeldi, a new Amazonian
Tree-Frog peculiar for its Breeding-habits. By Prof.
Dr. Emmi A. Gortp1, C.M.Z.8., Director of the Para
Museum
5. The Duke of Bedford’s Zoological Exploration in Eastern
~, Asia,—IIT. On Mammals obtained by Mr. M. P.
Anderson in the Philippine Islands. By Otprieip
MroMAS, HORS. EeZis.
Page
100
100
100
100
115
135
February 19, 1907. |
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of January 1907
Dr. C. I. Forsyth Major, F.Z.8. Exhibition of remains of a
Bear trounan Cay ernie Consicay nese crre eee enn
1. On English Domestic Cats. By R. I. Pocock, F.LS.,
F.Z.8., Superintendent of the Zoological Society’s
Gardens. (Plates VITI.—X.)
eee et eee ree eee estes eee ses eseseee
2. Report on Deaths occurring in the Society’s Menagerie
during 1906. By C. G. Srniemann, M.D., F.ZS.,
Pathologist to the Society
Pepe meee eee e enero reese reese sereeseese
3. On a peculiarly Abnormal Specimen of Turbot. By J. T.
CunnincHam, M.A., F.Z.S8. (Plate XI.)
eee eee reece se esece
4. On the Azygos Veins in the Mammalia. By Frank E.
Bepparp, M.A. (Oxon.), F.R.S., Prosector to the Society
5. Ideas on the Origin of Flight. By Dr. Baron Francis
Nopcsa
March 5, 1907.
The Hon. Walter Rothschild, M.P., F.Z.8. Exhibition of a
mountedyspecmmenior a) Govillaigetacac-acesa-i crise cee se
1. Descriptions of a new Species and two new Subspecies of
Antelopes and a new Sheep. By the Hon. Water
RopHscHiED Mims (Phas sHI ZR Stee ge oiiy. ds ceeaeaectias setae
2. On Elephant Remains from Crete, with Description of
Hlephas creticus, sp. n. By Dorornna M. A. Bare.
(Plates XII. & XIII.)
eee eee etme etree reser eons ee ess sess ees assoee
3. Zoological Results of the Third Tanganyika Expedition,
Page
143
168
174
181,
237
237
238
conducted by Dr. W. A. Cunnington, 1904—1905.— »
Report on the Polyzoa. By Cuarztes F. Rovusseer,
FRLMS. (Plates @kViGiXVs)08 te. Ae aelee tenes
V1
4. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905.—
Report on the Brachyurous Crustacea. By Wrixiam A.
Cunnineton, B.A., Ph.D., F.Z.S. (Plates XVI. &
XVIT.)
Ome mee w et eee eet eee reer reese een e te essere een eee ener ete seesu eee
5. On Two new Species of the African Genus Microchetus
belonging to the Collection of Oligocheta in the Museum
of Christiania. By Frank EH. Bepparp, M.A., F.R.S.,
Prosector to the Society
March 19, 1907.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of February 1907 .........
Mr. Herbert F. Standing. Notice of a Memoir on recently
discovered Subfossil Prosimiz from Madagascar
eee ee eeee
1. Descriptions of some New Species of Animal Parasites.
By L. W. Sampon, M.D., F.Z.8.
2. Descriptions of five New Species of Hemogregarines from
Snakes. By L. W. Sampon, M.D., F.Z.S., and C. G.
Seviemann, M.D., F.Z.8.
ii ii i i ii i i iii i aii ai ee ay
3. The Rudd Exploration of South Africa—VII. List of
Mammals obtained by Mr. Grant at Coguno, Inhambane.
By Ouprietp Tuomas, F.R.S., F.Z.S., and R. C.
Wrovuenton, F.Z.S.
April 9, 1907.
Mr. R. I. Pocock. Exhibition of a photograph and the skull
of a specimen! of Pallas’s Cat that had recently died in
the Menagerie, with general remarks on the species
1. On a small Collection of Fishes made in the Eastern
Watershed of the Transvaal by Capt. G. E. Bruce. By
G. A. Bounencrr, F.R.S., F.Z.S. (Plates XVIII. &
XIX.)
2. On the Winter Habits of the Greater Horseshoe, Rhino-
lophus ferrum-equinwm (Schreber), and other Cave-
haunting Bats. By T. A. Cowarp, F,Z.8. ...............
Page
258
277
281
28}
283
285
-. 2M)
307
vil
Page
3. Notes upon the Anatomy of aSpecies of Frog of the Genus
Megalophrys, with references to other Genera of
Batrachia. By Frank E. Bepparp, M.A., F.RS.,
Erosector toute; Society mn-reneeceeeeceai-sndassocse ae 324
4. Contributions to the Osteology of Birds.—Part IX.
Tyranni; Hirundines; Muscicape, Lani, and G'ymno-
nhines., ByaNVis Be Py Crani kh Anse eAelunS OCC mesa -ee ae 352
April 23, 1907.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of March 1907 ............ 319
Dr. A. Smith Woodward, F.R.S., F.Z.S. Exhibition of a
malformed antler of the Red Deer ....................000000% 380
Mr. R. I. Pocock. Exhibition of a model of the African
J Dlg lave at Go 0 Xs dee RP ARE irri, ocs ooceb accsucdscmoretoe’ 380
Mr. C. J. With. Notice of a Memoir entitled “* An Account
of the South-American Cheliferine in the Collections of
the British and Copenhagen Museums” .................. 380
1. The Ears as a Race-Character in the African Elephant.
BR Sb yD ORE ie said nis ain ie Sager n ete acne stele aeieotenbercine mses 380
2. The Duke of Bedford’s Zoological Exploration in Eastern
Asia.—IV. List of small Mammals from the Islands of
Saghalien and Hokkaido. By OuprreLtp Tuomas, F.R.S.,
F.Z.S. (With Appendix on the Cold-blooded Verte-
brates, by G. A. Bounencer, F.R.S., F.Z.8.)............... 404
3. On some new Species of Earthworms of the Family
Kudrilide, belonging to the Genera Polytoreutus, Neu-
mannielia, and Hminoscolex, from Mt. Ruwenzori. By
RANKS Hee BEDDARDs UV AY. THES user io: asee sda seeene aces A415
4. Some Notes on Hybrid Bears. By Henry SCHERREN,
BWA cgrtae reno s acide ices notte Na tlea har vst sls Vaccines snaee eee 431
hs
che
INIGIB BONING Lid
OF THE
CONTRIBUTORS,
With References to the several Articles contributed by each.
1907, pp. 1-446.
pp
Page
Baur, Purip Hernricn, B.A., F.Z.8.
On the “Bleating” or “ Drumming” of the Snipe
(Gallinage cochestis) .....0......ccsceuecee neces nen ene eenneceentees 12
Bare, Miss Dororuea M. A.
On Elephant Remains from Crete, with Description of
Elephas creticus, sp. n. (Plates XII. & XIII.) ...........: 238
Bepparp, Frank E., M.A., F.R.S., Prosector to the Society.
Contributions to the Knowledge of the Systematic
Arrangement and Anatomy of certain Genera and Species
Fok. Sofuaniarbaly eerste siesta os Leet vealed 30
On the Azygos Veins in the Mammalia ............-..++- 181
On Two new Species of the African Genus Microchetus
belonging to the Collection of Oligochzta in the Museum
Of Christiania. «acaeaesedsdesapoedsanseee Sasdacetemeesesen gene ee ee. SET
x
Bepparp, Frank E. (Continued.)
Notes upon the Anatomy of a Species of Frog of the
Genus Megalophrys, with references to other Genera of
IBEW A GX 0) 00 nar neat RMR ear MOE ator ons sAhdsea sec odoox
On some new Species of Earthworms of the Family
Hudrilide, belonging to the Genera Polytoreutus, New-
manniella, and Hminoscolex, from Mt. Ruwenzori
Bonuote, J. Lewis, M.A., F.LS., F.Z.8.
On a Collection of Mammals made by Dr. Vassal in
Aeninrermmy (@Plaibe ET, \? oo Seca) een: Seren eee Ut). 9
Bou.encer, Grorce AuBert, F.R.S., V.P.Z.S.
On a small Collection of Fishes made in the Eastern
Watershed of the Transvaal by Capt. G. E. Bruce.
(Plates XVIII. & XIX.)
Bee www eee eet were re eee e er esse se en sesene
On the Cold-blooded Vertebrates of the Islands of
Saghalien and Hokkaido. See THomas, OLDFIELD.
Catman, WittiAm Tuomas, D.Sc., F.Z.S., of the British
Museum (Natural History).
Notice of a paper on New or Rare Crustacea of the
Order Cumacea from the Collection of the Copenhagen
Museum: e222 eek) Re RY a ey I
CowarpD, THomas ALFRED, F.Z.S.
On the Winter Habits of the Greater Horseshoe,
Lhinolophus ferrum-equinum (Schreber), and other Cave-
hauintns Bats) 4h sees ckicclsncus dst aT ee ee
CUNNINGHAM, JosepH THomas, M.A., F.Z.S.
On a peculiarly Abnormal Specimen of Turbot.
(Plate XCM), 3:3... 385 das fede eae = gee ee
Page
415
307
100
312
x)
CuUNNINGTON, WiLLIAM ALFRED, B.A., Ph.D., F.Z.S.
Zoological Results of the Third Tanganyika Expedi-
tion, conducted by Dr. W. A. Cunnington, 1904-1905.
—Report on the Brachyurous Crustacea. (Plates XVI.
gC 2 I) es ie Re coro cn tL A oe ee
Duncan, F. Martin.
Cinematograph exhibition of Animals in the Society’s
Gardens and other Zoological Subjects
GoEtp1, Prof. Dr. Emi A., C.M.ZS.
On some new and insufficiently known Species of Mar-
moset Monkeys from the Amazonian Region
Description of Hyla resinifictria Goeldi, a new Ama-
zonian 'Tree-Frog peculiar for its Breeding-habits
Kenrick, Grorce Hamiuron, F.Z.S.
A List of Moths of the Family Pyralide collected by
A. EK. Pratt in British New Guinea in 1902-3, with
Descriptions of new Species. (Plates III. & IV.).........
Lankssrer, Sir Epwin Ray, K.C.B., M.A., D.Sc., LL.D.,
F.R.S., Director of the British Museum (Natural
History).
The Origin of the Lateral Horns of the Giraffe in
Foetal Life on the Area of the Parietal Bones ............
Parallel Hair-fringes and Colour-striping on the Face
of Fetal and Adult Giraffes. (Plate V.) ...............08.
On the Existence of Rudimentary Antlers in the Okapi.
(WHEE a iaiwin WAUL5) gaoesbuccedesbeccooocba64 oogucosboaedddiconde
Page
258
100
88
135
68
100
xil
Lyprexxer, Ricuarp, B.A., F.R.S., F.Z.S.
The Ears as a Race-Character in the African Elephant
Magor, Dr. C. I. Forsyru, F.Z.S.
Exhibition of remains of a Bear from a Cavern in
Corsica
Si) =heiesele.2)\2/.eie)s) aces ie ie)alaralei=)\sieleleleiele/e)\ele\s!a 0/6 o/slelle/«ielelalelsisisselalalelereienerehciniatctate
MircHEtt, Peter CHALMERS, M.A., DSe., LD. F.R:S.,
Secretary to the Society.
Report on the Additions to the Society’s Menagerie
during the months of November and December, 1906
eeceee
Report on the Additions to the Society’s Menagerie
during the month of January 1907
Cece er cece reese esse secs eces
Report on the Additions to the Society’s Menagerie
during the month of February 1907
Cece rsescesesvecesscssvesece
Report on the Additions to the Society’s Menagerie
during the month of March 1907
Seer cee errors so recess esses cene
Norcsa, Dr. Baron Francis.
Ideas on the Origin of Flight
Set eee ee sere secsccccre esse nsec
Pocock, Reervaup Iynzs, F.L.S., F.Z.8., Superintendent. of
the Gardens.
On English Domestic Cats. (Plates VIII.-X.).........
Exhibition of a photograph and the skull of a specimen
of Pallas’s Cat that had recently died in the Menagerie,
with general remarks on the species
ers eeeeercccoereesaovess
Exhibition of a model of the African Elephant
“« Jumbo ”
see ee oes e ses cescceresesesessovrersseesoceresocssceeesse essences
Page
380
143
281
379
223
143
299
Xi
Page
Pycrarr, WitLIAM Puaneg, F.Z.8., A.L.S8., of the British
Museum (Natural History).
Contributions to the Osteology of Birds.—Part IX.
Tyrannt; Hirundines; Muscicape, Lanti, and Gymno-
WIUATUCS:: sks. SMa SEN ets VN SOE RT Re TN REDE Ret Sk AR APL 352
Roruscuitp, The Hon. L. Watrer, M.P., D.Sc., Ph.D.,
E.Z8.
Exhibition of a mounted specimen of a Gorilla ...... 237
Descriptions of a new Species and two new Subspecies
Ot AMIE DES cinl Er SINEEO ooncoossdesosceonnuosoacss0e 6 Sead
RovussELET, CHARLES F., F.R.M.S.
Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904—1905,— Report
Gin Woe) IHolbywor,. (Ede NsGiss SUING ay DW ons easodoereenace: 250
Sampon, L. W., M.D., F.ZS.
Descriptions of some New Species of Animal Parasites. 282
Sampon, L. W., M.D., F.Z.8., and Seuiegmann, C. G., M.D.,
E.Z.8., Pathologist to the Society.
Descriptions of five New Species of Hemogregarines
EPROM SMA ROS Me eee ste arlene hole ieee eee lata se sas dala hela ae aun Sa: 283
ScHERREN, Henry, F.Z.S8.
Some sNoteston aly bridmbearcm eae era eee ee ee 431
XIV
Seviemann, C. G., M.D., F.Z.8., Pathologist to the Society.
Report on Deaths occurring in the Society’s Menagerie
Glpiaine? IMG ope seadessnnesdoasbenssbenouadde cose | codudasdnscssnnoc0s
Seriemann, C. G., M.D,, F.Z.8., Pathologist to the Society,
and Samson, L. W., M.D., F.Z.S.
Descriptions of five New Species of Hemogregarines
from Snakes..... Sr ROE NS IE la crc SO RE 3
Sranpine, Herpert F., M.Sc.
Notice of a Memoir on recently discovered Subfossil
TPrpogiraes tort INENORYSEING A 5 occccebacucco does apcocs00n030006
Tuomas, OLDFIELD, F.R.S., F.Z.S8.
Description of a new Monkey from the Ituri Forest.
(Plate VL.) 2.........ce0 seen ee cee tee tee ee ccee eters cents ectesecaeeeners
Exhibition of a collection of Mammals and Birds from
the Islands of Saghalien and Hokkaido .....................
The Duke of Bedford’s Zoological Exploration in
Eastern Asia.—III. On Mammals obtained by Mr. M.
P. Anderson in the Philippine Islands7..5..2.......0.......--
The Duke of Bedford’s Zoological Exploration in
Eastern Asia. —IV. List of small Mammals from the
Islands of Saghalien and Hokkaido. (With Appendix on
the Cold-blooded Vertebrates, by G. A. Bov.encrr,
TRISH AISE)) Scone vengspeusaaonocodsc0:-. he aise
‘Tomas, OLDFIELD, F.R.S., F.Z.S., and Wrovueuton, R. C.,
F.Z.S8.
The Rudd Exploration of South Africa,—VIT. List of
Mammals obtained by Mr. Grant at Coguno, Inhambane.
Page
168
283
281
bo
100
140
404
285
KV
Wirn, C. J., of Copenhagen.
Notice of a Memoir entitled “*An Account of the
South-American Cheliferine in the Collections of the
British and Copenhagen Museums tl a, se Ree ee
Woopwarp, Artuur Suiru, F.R.S., F.Z.S., Keeper of the
Geological Department of the British Museum.
Exhibition of a malformed antler of the Red Deer
eeeees
Wrovueuton, R. C., F.Z.S., and Tuomas, OLDFIELD, F.R.S.,
E.Z8.
The Rudd Exploration of South Africa—VII. List of
Mammals obtained by Mr. Grant at Coguno. Inhambane.
380
380
285
Plate
I.
VI.
TI. |
IV.)
We
VI.
Vii.
VILLI.
TN
Xe
MUL
ae
SS TUUL
IOV {|
Xv-}
XVI.
XVII.
XVIII.
XIX.
Proc.
LIST OF PLATES.
190i pps Elo:
Page
Cercopithecus dentt 2.0.2.0 ecu ce esse cc essere ete 2
NycEicebUs PYGMEUS 2.256. 0 cece eset et te ees 3
Pyralidee from British New Guinea .........+++..0-+> 68
Head of a Fetal Giraffe (half the natural size) ......-- 115
torn=tipsy orm Okerpietrs eet iarier so co vsatolee: (198
Young Ossicone or Horn of Okapi..........-+. 200000 Fi
Blotched Tabby, Felis catus ......--..e eset ]
Striped Tabby, Felis torquata ..........00 sees beeen , 143
Agriotypes of the Striped Tabby
Nbnormal VouneMurbot 2: 4. oti et 174
Elephant Remains from Crete........00.+.s.-eee eres 238
Tanganyika Polyzoa ........--.0-ee eee eer s teens 250
1. Potamen (Potamonautes) orbitospinus. 2-7. Antennal
Region of various Potamonidee .......-..--+++++> A
1&3. Potamon (Potumonautes) platynotus. 2 & 4. Platy- | a
thelphusa conculeata. 5 & 6. Platythelphusa maculata
1. Barbus bructt. 2. B. sector... 00.0. cee e nese sence | «
Varicorhinus brucit . 6.0... eect ees: \
Foou. Sov.— 1907. h
Vial) ibn cca
ae: en ‘lib Res dept sgn
yeti! Mirksiies
me
ae, Bi oe ie
re Ey atten
eect pest
| “hh pdt A 6 oO vabinalatiagl
ff. ee Cee sn a
Das) ot Mee ee
C2 bo
LIS? OF TEXT-FIGURES.
1907, pp. 1-446.
. Palatal view of skull of Nycticebus pygmeus.. cc... ceveen.
. Lateral view of skull of Mycticebus pygmeus... 0... cece eee
. a. Snipe bleating, showing characteristic position. 6, Formation
Oli tallbaszordimeanilw ibe! deinetic: hime re teenie
. Varieties of outer tail-feathers of Gallinago cwlestis ........
. a, Dorsal view of musculature of tail of Snipe. 6. Ventral
view of musculature of tail of Snipe ...............00008
. A. Half of tail of Gallinago celestis from without inwards
left to right. B. Half of tail of G. delicata from without
inwards left to right. C. Half of tail of G. gullinula ......
. A. Section of two rami of a feather showing interlocking of
distal and proximal radii. B. Ramus of Gallinago celestis,
showing proximal and distal rows of radii. C. Distal radius
of G. celestis. D. Proximal radius. E, Distal radius of
middle-tail feathers of G. calestis.........0. cc eves seeeees
8. A. Half of tail of Gailinge major from without inwards left
to right. B. Half of tail of G. nobilis from without inwards
left to might ©: Elalfion tailot Ge stenw@ 55.0405
9, a. Tail of Gallinago solitaria. 6. Tail of G. megala ........
10. Lung of Chameleon verrucosus, entire ........000 0. eevee
ie Wune ot Chameleonidilepis entire wens. + ssc ls. eee
12. Lung of Chameleon parvilobus, opened longitudinally ........
13. Lung of Chameleolis, opened longitudinally ................
14, A. Stomach and pancreas of Chameleolis. B. Dorsal aorta
ATG MAMIE Deine ENE No LE BWasbaeoeseoedul si cow we awe
l5eMkgua sconcowdes Weadand meck sai j4. 2 oes
16, A portion of the trachea, the bronchi, and the upper parts of
the lungs of Heloderma, fvom the dorsal side ..............
Page
5
5
16
18
i)
bo
23
poco
“I
be te woe el
= S
bo bo
Co bs
ASS
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~t
36.
37.
39.
40.
4l.
Oo
KX
Page
. Pancreas, gall-bladder, &c., of Varanus niloticus, to show the
course of the bile-ducts .......... a haa pial ele Sata ea GI tice 64
IDO) LE [Arie WIS CMO OTOCHOHIES. ooo co ne@adounsoobannebasor 65
PUG SOL WAKAWUS GIUSEUS). .. «5. o)eeMeenne anon oe eae eae 66
Mbackeviewsothead Olu VVd as C/(OUtes ime aetna 89
Mehack views of headuot Midas 7 /Ucentcrassmr ere eee 6 Gil
A Backvie, of lead of Mudasigniscoventea 1. eee seer 93
milileadsote Midas nye ator |. sateen 94
. Lateral view of the skull and lower jaw of a very youny
Giralle. sis wane ero anes eee © Pongtie aeep aan 101
, Lateral view of the skull of a Giraffe, about two-thirds grown. 102
26. View from above of the fronto-parietal region of the skull of a
WEIAP ROUTES EDN RNIE Rea gaoo suo dando ds cl Gon OO HdMmaD eo bG oA. 102
. View from above of the fronto-parietal region of the skull of an
duane tute MO Ka pi wis aveieraagueva tse, asa tor occ usteenemetewerstate terete suelo seecett 103
28. Sagittal section through the bony tissue of the ossicone and
the roof of the skull of a very young Giratfe .............. 104
. Copy of the drawing of a sagittal section through the ossicone
and the roof of the skull of a newly-born Giraffe, published
by the late Sir Richard Owen in the Trans. Zool. Soc. 1840.. 104
. Drawing of the right side of the head of the fcetal Giraffe
described in the text .......:....... BASES. 6 SOR wae 106
Uhesleic shormesoiethe) tcetalmGntalcmnn tyne irre eo nian ot 107
. Diagram to show the flattened plate-like form smi the
orientation of the horns ef the foetal Giraffe .............. 108
33. Skull of Giraffe....showing the directions in which the
Slut welsh CULM: fraatre fea Deocaerensrn catnenewes cisia jes Rerpiere Meets 110
. View, looking forwards, of the skull cut in the direction of
ivory 424 a WORE SECM SE CMbiclke asa wa occ onesies codahblds Dak dil
. Drawing, of the natural size, of a young foetal skull of a
(CHIPTEHE Reta ae eS eR amen ae aes aR ake cha Bis Hines ers Se ey nae 112
Diagram to show the various positions on the frontal bone at
which the bony horn-cores of the Cavicorn and Cervine
Tuminants may take them erowth\..- s-)) e eee reece - 114
Left side of the head of the foetal Giraffe described, showing
colour-stripes on the snout and above and below the eye.... 115
. Section across three dark-coloured bands above the eye of the
foetal Giraffe, showing three longitudinal furrows or de-
pressions in transverse section corresponding to the dark
WHC mc oeadescetesero vos cs ado essen seovsdbecosagmaaE 116
Section of the skin of the frontal region of the foetal Giraffe,
showing threesizes Of Waits Terr miele elaine ela aie-/ ( Lily
Surface-view of banded “pelage”’ of the frontal region of foetal
Giraffe, showing the convergence of the hairs at the three
longitudinal Tiana Oi Gawd ayyOCRMNNOS 5 ocdococsod.essccnce 118
An enlarged drawing of a single hair of average size ofr om the
frontal region of an adult Giraffe, showing the ‘dark pigmented
free extremity and the opaque white lower DOAN “S sueuees 119
42.
y
; Colour-striping on the face of a Somali-land Beant . chain teegiiths 124
. The same specimen as that shown in text-fig. 47, with colour
XX1
Page
Colour-striping on the face of an adult male Giraffe from
Kordofanvirraa i) 4. ae ey Sepak st, Atl areecare Hiaiith 120
. Colour-striping on the face of an adult female Giraffe from
ond ofan ree Oy Soy Pay hou edo ceaneapebisieh aisha bis 120
. Colour-striping on the Anas al Giraffa ae dalis cottont
from: Mitz HleonsUWeanday icy anne ‘ RU avant ata Ay
: Colour-striping on th face and muzzle ofa a Thaiernsel Giraffe... 122
. The same specimen as that shown in text-fig. 45, with colour
OMMUETERE: Aye Ye). CON aeaee Bvt capay betas Fe Reet ee ile
Onna GS Pea ol I SRS. 8 Nk CRY EN) ecu NURS ea ie HERR 124
. Drawing of a fore and aft section through the tip of the ossicone
of amt adults Okapi | Ase A) eects: & skaters byevenateverenne as tasers acer 126
5), Rudimentary free ossicone of hemispherical shape from the
skin overlying the frontal bossed region of the skull of an
Okapitofethe broud-siculleditype) Wank aes see aces niet 128
51. Section of the ossicusp drawn in text-fig. 50, to show the
In{coOmlpleter ossilie sti Om mej vecvalss 9 -)-Naate ene enekeielsy siotaet ate cist etele 128
52. Bisected horn-tip of the Edinburgh Okapi...............055 13L
53. Section of half of the same ‘Waecal OMM=bip Ns. 1 « 131
64. Drawing of the right ossicone of the Okapi brought by Capt.
Boyd Alexander “Fhonin the Welle River ...... Maint: tovetoers 154
55. Section through the tip of the same specimen to show the
incomplete trabeculated Ossification’............---.s++5+- 134
DGwerylal resinafict) Uy Mule wit acts ei qumGics shiseier ni eke a aye 136
is Jalil Tay vonee (Cheerenarnes@) a8 s auoercokceseeoun aud 137
58. Breeding-basin of Myla resinifictrix: (A) side view, and (B)
OOM) (Chien TMG) 2 oo oe Soe Ue bo od od 5544506 voueS a 158
59. Breeding-basin of Hyla resinifictria, seen from above ..... oe ltd
60. Diagram of the Pattern of the Blotched Tabby (Felis catus) .. 154
61. Aorta of Tiger, showing several aneurysms ............ So te. LEG
G2wAZyeos vems.ol Cenvicapia Donota. ses ee ass ia- Seetatiel to atniolels 186
Gay Aayeos veins of GarellaneiGhone wniiiiess “ii. - its lreers iets sa 187
64. Azygos veins of Phacocherus ethiopicus.... 0.0.0.0 eceee snes 189
65. Azyvos veins of Tragulus menwnnd). 0.0605. o ce tweens 191
66. Hyrax capensis; two different arrangements of azygos system. 194
Gi Azye0s veins Of Lrvnaceus agus) J... o-tiriansr ais 196
GSH AZygosiOn Ge ANCrOcuba, Wn acto a twle SIP) « di-siewinle: «Weyl tae 199
69. Azygos of Crossarchus ohscurus .......+... siiaeval avec eens .. 200
70. Azygos veins of Halmaturus bennett .........0.0..20 ee cease 203
71. Azygos veins of Phascolomys mitchellt ........0.ssecvevees 204
(Pe ENA) veils Ol JESUP INGEN. 1) 5 oo BD bens au eno uo a Seo 211
73. Lateral view of the thoracic region of newly born Myopotamus
COU PU wa « sohansrent eee mtcR Neier dee) a Seater et Nelen ee tiedeine caeF. Meena a Sr alsa 214
74. Hind limb at Diangimhodor Peas ee te eaeen ee eng Samia 4) DO
“ow blind limbs on Rhamprorymenuss Wiz. aeiee atest! ths eis 225
AGsullingdslim'a Ob Pia acany ism ose Rene ars ae ait retasts 226
XXIl
Page
(i odind timbyot Wyetodactylus 0)... ee. 227
(37 Hindllimbsok Hipposiderus —)......). eee net ee 30
(OmEindlimbrot Ornichomiumus sso 0. 2) se. ee eee eye, On
80. Hind limbs of Dipus (left) and Alactaya (right) ............ 232
ily [altos Whirl) ore UA ACA COED Jobb bobcs cua bagdsonassdansen 233
82. Hypothetical reconstruction of a running “ Pro-Avis” ...... 235
85. Right mandibular ramus of Elephas antiquus (?) bearing two
lower:molars’ Zan. ¢. aaa Mee es oct eee 245
84, Platythelphusa armata, large male ...........00-000ceecaes 269
80: Micracketusicollctiy a nee ee ee, ee ee 278
BONUMicrochelusiZulicnsisi ie. eh anes eaten 280
Oe Helis maniiean scat are eee 6 cB rar 55 300
88. Skull of Felis manu!, view from above .............0000008 304.
Go” Skullloterelsunanulssideicw eae ee ee 305
SO Barbus polylends: 1a a Geay Ce amen cerns 2 ern ots RE 308
Oy Gonbusrelephuntis® hoe). (en eee ee oe OO,
92, Palmar surface of hand and foot of Me Halonney VS WOBHED eens o 326
93. Part of dorsal musculature of Wegalophrys nasuta............ 302
94, Some of the dorsal muscles of Rana gunpyt ...............- 333
95. A dissection of Pelobates fuscus, to show large cesophageal
sheet ot thestransyerselich see nnn e cn 304
96. Some of the dorsal muscles of Ceratophrys orndta .......... 336
ine klvoid tof Megalophnys nasutcmne teen 341
98. The cesophageal sheet of transversalis muscle in Megalophrys
OSES 65 85 ce MONT Sb B1e CRE RE RT En ete 346
99, Liver and adjacent viscera of Rana guppyt ....... cee cee ees 349
OOO viductotMegalopinysacasuta mee ee ee ee 350
101. Lateral aspects of the skulls of :—a. Gymnorhina organicum.
b. Laniarius poliocephalus. e. Sayornts cineracea ........ 355
. Ventral aspects of the skulls of :—a. Artamus leucogaster.
b. Sayornis cineracea. cc. Terpsiphone. 4d. Pitta baudi.
e. Vireolanius leucotes. {. Vireo olivaced ...............- 362
103. Ventral aspects of the skulls of :—a. Zanius collurio. b. Bas a-
disea raygiana. c. Gymnorhina organicum................ 363
104. Phylogenetic tree indicating the probable relationships of the
Passeres Oligomyodii and Diacromyodii .................. 377
105, Head of the Addo Bush, or East Cape Elephant (Hlephas
G/PUCANUS ICAPENSIS) «2. e714. Se ee, 383
106. Head of Female West Cape Blephant (Elephas africanus
SOU EES EOE AS DORE EM RA. fs Acs 5 63-0. .n:6.0 080-Ao RE 385
107. Head of Male West Cape Elephant (lephas africanus
OUR ES) PRE AIAN Pee TR AMADA IN cc rh cin Gobioes Ad Chere 386
108. Head of Male Mashonaland Elephant (Elephas africanus
BELOUSL) a, SN, Fake Ie SORES ARE eee ree tere emt th 387
109. Right Ear of Male Mashonaland Elephant (Elephas africanus
SCLOUSD La ee ha Shia ence La ee ane: 388
110. Head of Male Elephant from Swaziland .................. 389
111. Right Ear of the Congo Elephant (Zlephas africanus cyclotis?). 390
XX1ll
Page
112. Head of a Male Elephant from North-west Rhodesia ........ 391
113. Head of Male Elephant from Fort Manning, N.E. Rhodesia
(Hlephas afiracanus knochenhawert?) .....5..g..-+-s02++-- 392
114. Head of Male Elephant from the Aberdare Mountains, British
Wast, Autica rants: os s)s ac naeonrame Renan rience a oth sald ~- 393
115. Head of Male Elephant from the Lake Rudolf District (Zlephas
AFTICONUS COVENGISHE)) io an cheEd SOTA ames eS roca, 8 ett 394
116. Head of the “Queen’s Elephant,” an immature Male Sudan
Hlephant, (sage ..cc ed 5) ete eR eer ae eerie. 396
117. Right Ear of a Male Sudan Elephant (Llephas africanus
OCU OLISI) Ys, she sie eons So Whey lag sno aah ees ETA RI La NRE RS = 397
118. Right Ear of the North Somali Elephant (Zlephas africanus
OUUCAIST Nin ae sales os osle sah sietanoneiearaiat eGR aa ele euSASRDs ean heen 398
119, Head of “ Jumbo,” the male West Sudan Elephant formerly
belongineytonbhne: Socte tyes tae ieee sie eerie 400
120. Front view of the Skull of the Sudan Elephant (Llephas
A ILC ANUS! OMYOLUS) aise Mea sbeu Masses 2 isn, PARENTS © eich eta oe 40]
121. Front view of the Skull of the Albert Nyanza Elephant
(Hlenhasyajnicaniusialbetersts) ei een eel ae eae 402
122. Spermathecal sac of Polytoreutus ruwenzorii ......- +02 eee 418
128. Yerminal male organs of Polytoreutus granti .......0. 2.005. 42]
124, Spermathecal sac of Polytoreutus grant? ...............44. 422
125, Spermathecal apparatus of Newmanniella ruwenzori.......... 426
126. Ventral view of Eminoscoler ruwenzorat ... ........+-.-+- 498
127. Terminal male organs of Lminoscoler ruwenzortt ...........5 430
LIST OF NEW GENERIC TERMS
PROPOSED IN THE PRESENT VOLUME (pp. 1-446).
Monodonta (Insecta)...... 69 | Tipuliforma (Insecta) ......... ..
PROCEEDINGS -
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
| ZOOLOGICAL SOCIETY
OF LONDON “eel
1907.
Paces 1-236.
CONTAINING PAPERS READ IN
- JANUARY anv FEBRUARY.
JUNE 1907.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDON :.
MESSRS. LONGMANS, GREEN, AND CoO.,
; PATERNOSTER-ROW,
ae —Price Tiuelve Shilings] ERY
aGenal Musee
LIST OF: CONTENTS:
1907, pp. 1-236.
January 15, 1907.
Page
The Secretary. Report on the Additions to the Society's Menagerie during the months of
November and December, 1906 .......- eee cece cece ee erence Sistele se ueuvenne wistnicuste 1:
Mr. Oldfield Thomas, F.R.S., F.Z.S. Description of a new Monkey from He Ituri
Forest. (Plate 1.) 2... cect eee cect ce cece ees cece ec cece eas essne eens stones 2
1. On a Collection of Mammals made by Dr. Yassal in Annam. By J. Laws Bonnorg,
M.A., F.LS., F.Z.8. (Plate II.) ..... PEPE S Arias) ah eee heen Lay AC Demo ri. & 3
2. On the “ Bleating” or “ Drumming ” of the Snipe (Gallinago cwlestis). By P. H. Baur,
TRAV SAHIN nia epee stalcde 10 gietei is en lace eh amie aoe Re teal emieieieh ty. Uo tate datevale 6 whe lavas feoe Bian iiee ls
3. Contributions to the Knowledge of the Systematic Arrangement and Anatomy of certain —
Genera and Species of Squamata. By Franz EB. Bepparp, M.A., F.R.S., Prosector to
bE OCI by sa < vere ia allo eis cie >'» Sie vetets sie eh ee ae a eemar ea aisle Notcetarets A Hatem 14 35
4. A List of Moths of the Family Pyralide collected by A. EH. Pratt in British New Guinea
in 1902-3, with Descriptions of new Species. By Gnorczk H. Kenrick, ¥.Z8.
(Pints WUT, DV) Fie vies otaemie esanclse ale een retrere Pstarete AE Fy Ae a ES oh 68
5. On some new and insufliciently known Species of Marmoset Monkeys from the
Amazonian Region. By Prof, Dr. Bui A. Gorrnr, C.M.ZS., Director of the Para
Museum 2.0.6 cece eee te ee cece reece eet ee tnt et ene cnet eee cette es Arata 88
February 5, 1907. ;
Myr. F. Martin Duncan. Cinematograph exhibition of Animals in the Society’s Gardens
and other Zodloridal Subjects. ceo POH es ihe wo aera 2 wiahinten sian ate acsiate siete tate tetele 100.
Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition of a collection of Mammals and Birds
from the Islands of Saghalien and Hokkaido ........6..cseeeeeee SR sna c lee SOU
Contents continued on page 3 of Wrapper.
THE ZOOLOGICAL SOCIETY OF LONDON.
eee
Tuts Society was founded in 1826 by Sir Sramrorp Rarrtes,
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vAG,, M1). | s
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ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINESS.
(AT 3 HANOVER SQUARE, W.)
1907.
TuEspay, JANUARY 15 Turspay, May .... 7 and 28
¥y Feprvary 5 and 19 if JUNE ees
és March .. 5 «,- 19 56 November 12 and 26
- AUER oy SN oy 28} ., DecremBer 10
The Chair will be taken at half-past Eight o'clock in the Evening
precisely.
LIST OF THE PUBLICATIONS
OF THE
ZOOLOGICAL SOCIETY OF LONDON,
Tue scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and “ Transactions,” in quarto.
According to the present arrangements, the ‘‘ Proceedings”’
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the ‘‘ Proceedings” by
the Committee of Publication. A large number of coloured
plates and engravings are issued in the volumes of the
“ Proceedings,” to illustrate the new or otherwise remark-
able species of animals describedin them. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society’s Gardens are often given.
The “ Proceedings” for each year are issued in four parts,
on the first of the months of June, August, October, and
April, the part published in April completing the volume
for the last half of the preceding year. From January 1901
they have been issued as two half-yearly volumes.
The ‘‘ Transactions” contain such of the more important
communications made to the scientific meetings of the Society
as, on account of the nature of the plates required to illustrate
them, are better adapted for publication in the quarto form.
They are issued at irregular intervals.
Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea before the day of the Anni-
versary Meeting in each year, are entitled to receive the
Society’s Publications for the year. They are likewise
entitled to purchase the Publications of the Society at 25 per
cent. less than the price charged for them to the Public. A
further reduction of 25 per cent. is made upon purchases of
Publications issued prior to 1881, if they exceed the value of
five pounds.
Fellows also have the privilege of subscribing to the
Annual Volume of the Zoological Record for a sum of 30s.
(which includes cost of delivery), payable on the Ist July
in each year; but this privilege is forfeited unless the
subscription be paid before the 1st of December following.
The following is a complete list of the publications of the
Society already issued.
[ June, 1907. ]
TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.
4to. 16 vols. and Index. Enice te Price to the
ellows. Public.
Vol. _I., containing 59 Plates.... (1833-85) .... £3 13 6.... £4 18 Of
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Vol. XI., containing 97 SBIR: 3 (U880=85) a OD Oa al eG uO
i X11, yy | OD 2 yy? oo (esGH80) sogn O 8 O; i 4
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PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF
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9 i)
PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
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= WUE, Ike), op 4s. Gd. .. 6s. $9 XI. 1848. he 4s. 6d. ... 6s.
Pe LValeoG. 7H 4s. 6d. .. 6s. 5 ue aise: = 4s, 6d. .. 6s.
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{ SOOWE IS, 9 4 Gs: em 1ll 6 2 2 Ot
Index 1848-1860. el a Gch 6s.
t Out of print.
* In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is
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separate papers at about one-fourth their published price,
PROCEEDINGS
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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Highth Edition.) 8vo.
1883. Cloth, 4s. 6d.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Ninth Edition.) 8vo.
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Catalogue of the Library of the Zoological Society of London.
(Fifth Edition.) 8vo. 1902. Cloth, 6s.; Paper, 5s.
These publications may be obtained at the Socrery’s OFFIce
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THE ZOOLOGICAL RECORD.
——-05900——.
THE object of the Zootogican Record is to give, by means of an
annual Volume, complete lists of the Works and Publications
relating to Zoology in all its branches that have appeared during
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parts of the globe, and thus to form a repertory that will retain its
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The ‘ Zoological Record’ is published by the Society at the price
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The divisions of the ‘Zoological Record,’ commencing with
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SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.
At present each Volume of the Zoonocicat Rucorp consists of
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in paper covers, stitched and lettered.
The following are the Divisions and their net prices, viz. :—
Se ile
List of abbreviations of journals, ete. 2a)
Special Records, viz. :—
I. General Subjects .. 26
II. Mammalia RB 8
III. Aves acted a riecn ma 6 0
IY. Reptilia and Batrachia.. Ab
V. Pisces 2 6
VI. Tunicata ue
VII. Mollusea 4 0
VIII. Brachiopoda .. ile
IX. Bryozoa I ©
X. Crustacea oy 8
XI. Arachnida 4)
XII. Myriopoda pe
XIII. Insecta .. 12 0
XIY. Echinoderma a) 6
XGVEN Wiermes >. oy
XVI. Coelenterata . Then 8
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XVIII. Protozoa 2)
Index of new names of genera and subgenera 2)
On receipt of the price any Division will be forwarded as soon
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These separate Divisions can be obtained trom the Zoological
Society, 8 Hanover Square, London, and also from Messrs. Fried-
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Orders should be made payable to “The Zoological Society,” and
crossed ‘“* Drummond’s,”
P. CHALMERS MITCHELL, M.A., D.Sc., LL.D., F.R.S.,
Secretary.
June, 1907.
ZOOLOGICAL Soctety or Lonpon,
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LIST OF VOLUMES or raz ‘ZOOLOGICAL RECORD,’
The Record of Zoological Literature, 1864-1866, Vols. 1.-111.,
and 1868, Vol. v. Edited by Anserr C. L. G. Ginrner, M.A.,
M.D., Ph.D., F.Z.8., &e. Price 10s. each Volume. Net. (1867,
Vol. rv., supplied with sets only.)
The Record of Zoological Literature, 1869, Vol. vr. Hdited by
Atsert C. L. G. Gintner, M.A., M.D., Ph.D., F.R.S., F.2Z.S., &c.
London, 1870. Price 30s.
The Zoological Record for 1870-1872, Vols. vir.tx. Hdited
by Atrrep Newroy, M.A., F.R.S., F.L.S., V.P.Z.8., &e. Price 10s.
each Volume. Net.
The Zoological Record for 1873-1883, Vols. x._xx. Edited by
Epwarp Canpwett Rys, F.Z.8., M.E.S. Price 10s. each Volume. Net.
The Zoological Record for 1884, 1885, Vols. xx1., xx11. Edited
by F. Jerrrey Bett, M.A. Price 10s. each Volume. Net.
The Zoological Record for 1886-1890, Vols. xx1i1.-xxvit.
Edited by Frank E. Brepparp, M.A., F.Z.8. Price 10s. each
Volume. Net.
The Zoological Record for 1891-1900, Vols. xxviII.—xxxvil.
Edited by D. Saare, M.A., F.R.S., F.Z.8., &c. Price 10s. each
Volume. Net.
The Zoological Record, Volume the Thirty-eighth; being
Records of Zoological Literature relating chiefly to the year 1901.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
Alice L. Embleton, EK. R. Sykes, E. A. Smith, 8. Pace, Albert
Brown, D. Sharp, F. A. Bather, and E. A. Minchin. Fdited (for
the Zoological Society of London) by Davin Suarp, M.A., F.R.S.
F.Z.8., &c. London, 1902. Price 30s.
The Zoological Record, Volume the Thirty-ninth ; being Records
of Zoological Literature relating chiefly to the year 1902, By
D. Sharp, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, W. T.
Calman, E. R. Sykes, E. A. Smith, Alice L. Embleton, F. A. Bather,
E. A. Minchin, and H. M. Woodcock. Edited (for the Zoological
Society of London) by Davin Suarp, M.A., F.RS., F.Z.8., &c.
London, 1903. Price 30s.
The Zoological Record, Volume the Fortieth; being Records of
Zoological Literature relating chiefly to the year 1903. By D.
Sharp, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, W. T.
Calman, E. R. Sykes, E. A. Smith, Alice L. Embleton, F. A. Bather,
E. A. Minchin, and H. M. Woodcock. Edited (for the Zoological
Society of London) by Davip Suarr, M.A., F.R.S., F.Z.S8., &e.
London, 1904. Price 30s.
The Zoological Record, Volume the Forty-first; being Records
of Zoological Literature relating chiefly to the year 1904. By D.
Sharp, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, W. 1.
Calman, E. R. Sykes, E. A. Smith, Alice L. Embleton, F. Silvestri,
EK. Simon, F. A. Bather, W. Woodland, and H. M. Woodcock.
Edited (for the Zoological Society of London) by Davin Suarp,
M.A., F.R.S., F.Z.S., &. London, 1905. Price 40s.
The Zoological Record, Volume the Forty-second ; being
Records of Zoological Literature relating chiefly to the year 1905.
By D. Sharp, R. Lydekker, R. Bowdler Sharpe, Igerna B. J. Sollas,
W. T. Calman, EK. R. Sykes, E. A. Smith, Helen P. Kemp,
F, Silvestri, A. 8. Hirst, Margaret Grant, Cora B. Sanders, E. A.
Minchin, and H. M. Woodcock. Ldited (for the Zoological Society
of London) by Davin Sarr, M.A., F.R.S., F.Z.8., &. London,
1906. Price 40s.
Index Zoologicus. An alphabetical list of names of genera
and subgenera proposed for use in Zoology, as recorded in the
Zoological Record, 1880-1900; together with other names not
included in the ‘ Nomenclator Zoologicus’ of S. H. Scudder. Com-
piled (for the Zoological Society of London) by CHarztus Owen
WarterHouse and edited by Davin Suarp, Editor of the Zoological
Record. London, 1902. Price to Fellows, 18s.; price to the
public, 20s.
These publications may be obtained at the Socrety’s OFFICE
(3 Hanover Square, W.).
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
Or THE
ZOOLOGICAL SOCIETY OF LONDON.
(January to April, 1907.)
January 15, 1907.
Dr. J. Rosz Braprorp, F.R.S., Vice-President,
in the Chair.
The Secretary vead the following report on the additions that
had been made to the Society’s Menagerie in November and
December 1906 :—
The registered additions to the Society’s Menagerie during the
month of November were 173 in number. Of these 105 were
acquired by presentation and 23 by purchase, 35 were received on
deposit, 3 in exchange, and 7 were born in the Gardens. The
total number of departures during the same period, by death and
removals, was 177.
Amongst the additions special attention may be directed to :—
An adult male Mandrill (Papio maimon), the first full-sized
example of this species exhibited in the Gardens, deposited on
Noy. 30th.
A young female Hippopotamus (Hippopotamus amphibius) trom
the Niger, purchased on Novy. Ist.
A Persian Stag (Cervus maral), presented on Nov. 13th by
Mr. Carl Hagenbeck.
A Kashmir Stag (Cervus cashnuriensis), presented on Noy. 22nd
by H.G. the Duke of Bedford, K.G., P.Z.8.
A Collection of 47 Birds, including, among other interesting
species, a Toucan (Awlacorhamphus sulcatus), new to the Collection,
Proc. Zoou. Soc.—1907, No. I. 1
2 ON A NEW MONKEY FROM THE ITURI FoREST. [Jan. 15,
anda Sun-Bittern (Hurypyga helias) from Venezuela, presented
on Nov. 27th by Capt. A. Pam, F.Z.8.
The registered additions to the Society’s Menagerie during the
month of December were 150 in number. Of these 67 were
acquired by presentation and 16 by purchase, 61 were received on
deposit, 2 in exchange, and 4 were born in the Gardens. The
total number of departures during the same period, by death and
removals, was 207.
Amongst the additions special attention may be directed to :—
A pair of Siberian Dholes (Cuon alpinus) from Thian Shan,
received in exchange on Dec. 2nd, new to the Collection.
A Cape Hunting-Dog (Lycaon pictus) from South Africa, pur-
chased on Dee. Ist.
An Addax Antelope (Addax naso-maculatus) from North Africa,
presented by H.G. the Duke of Bedford, K.G., P.Z.S., on Dec. 18th.
A Bubaline Hartebeest (Alcelaphus bubalinus), and a hybrid
between Pére David’s Deer (Hlaphurus davidianus) and the Red
Deer (Cervus elaphus), deposited on Dee. 29th.
A new Monkey from the Ituri Forest.
Mr. Oldfield Thomas, F.R.S., F.Z.S., exhibited a Monkey which
had been obtained in the Ituri Forest, Upper Congo, during the
recent Ruwenzori Expedition, and gave the following description
of it :—
CERCOPITHECUS DENTI Thos.* (Plate I.)
Abstr. P. Z.S. 1907, p. 1 (Jan. 22, 1907).
A member of the campbelli-mona group, but not darkened on
the posterior back and hind limbs, and with a very sharply con-
trasted white belly.
Upper surface of head and neck olive-grey, the usual light frontal
baud present but not conspicuous. Back dark grizzled chestnut-
brown (nearest to “burnt-umber” of Ridgway); colour of rump
not darker, but, on the contrary, passing gradually into the
paler tone of the hips and hind legs. Under surface from chin
to anus, and inner sides of limbs to wrists and ankles, clear
creamy-white, very sharply defined from the darker colour, not
only on the limbs, as in campbelli and others, but also along the
flanks, where the white rises nearly halfway up the lateral aspect
of the animal. Ears with short yellowish tufts rising from their
inner surfaces. Outer sides of fore limbs deep black from
elbows. Hind limbs grizzled yellowish olive, lighter than the
back, down to and including the ankles, the metatarsals and toes
black. Tail indistinctly blackish above at base, then dull greyish
white for two-thirds its length, darkening again to black on its
terminal third.
* [The complete account of the new species described in this communication
appears here ; but since the name and preliminary diagnosis were published in the
‘Abstract,’ the species is distinguished by the name being underlined.—Eprror. |
SZ SIO yvealk 1.
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1907. | ON MAMMALS FROM ANNAM. 3
Dimensions of the type, measured in the flesh :—
Head and body 501 mm.; tail 850; hind foot 155; ear 40.
Skull— greatest length 105 mm., basal length 75; breadth of
brain-case 55; length of upper cheek-tooth series 23.
Hab. Ituri River, between Mawambi and Avakubi, Upper
Congo ; alt. 8000’.
Type. Adult male. B.M. no, 7.1.2.1. Original number 184,
Collected 23 October, 1906, by R. HE. Dent.
This handsome Monkey 1s most nearly allied to the W. African
C. campbelli, but differs by its grizzled olive-yellowish instead of
black hind limbs, the absence of black on its posterior back, its
more or less greyish-white tail, and by the high and sharply
defined line separating the colours of the flanks and belly .
The following papers were read :—
to}
1. On a Collection of Mammals made by Dr. Vassal in
Annam. By J. Lewis Boynore, M.A., F.L.S., F.Z.8.*
[Received November 16, 1906. ]
(Plate IL.7 and Text-figures 1, 2.)
The British Museum has recently acquired a most interesting
set of Mammals from Annam, collected by Dr. Vassal. The
collection contains examples of some twenty-five species, of which
five are new to science, while several of the others add consider-
ably to our knowledge (still very limited) of the fauna of the
district.
Since the collections made by MM. Pierre and Mouhot over
half a century ago, practically no fresh material has reached
Europe from that locality. As would therefore be expected,
many of the forms are undescribed, and there is little doubt that
with further material many of the forms at present included
under existing names will prove to be subspecifically distinct.
The collection is, perhaps, too small for any generalisation on the
fauna of Annam, but its affinities seem if anything to tend towards
China rather than the Matay Peninsula, and it is especially note-
worthy that it differs considerably from the fauna of Siam. Lest
I am misunderstood, I may as well point out that by “fauna” I
am not referring to the presence or absence of certain genera,
but rather to the fact that the local forms of widely spread
species approximate rather to the Chinese than to the Malayan.
To give some examples:—The Porcupine is Anderson’s Hystria
yunnanensis, not H, groter from the Peninsula. The Petaurista
is Anderson’s P. yunnanensis, and not P. lylei, mihi, from Siam.
The new Zupaia described has its affinities with 7. chinensis and
* [The complete account of the new species described in this communication
appears here; but since the names and preliminary diagnoses were published in the
* Abstract,’ such species are distinguished by the name being underlined.—Ep1rTor. |
+ For explanation of the Plate, see p. 11.
1*
4 MR. J. L. BONHOTE ON [Jan. 15,
not with 7’. ferruginea. On the other hand, the Paradoxurus is
apparently identical with a form described by me from the
Peninsula. Another point of interest as showing a probable
double origin for this fauna, is in the occurrence at the same place
of two subspecies of Sciwrus macelellandi—one, S. m. rodolphi
A. M.-E., showing very obvious affinities with S. m. barbei of
the Peninsula; the other, S. m. maritiémus mihi, which is in-
distinguishable from the type, which came from China. It must,
however, be remembered that this last is only represented by a
single skin, and it might possibly have been brought down on a
ship and escaped.
Lastly, attention may be called to a new species of Vycticebus,
which is in many respects intermediate in its characters between
Nycticebus and Loris.
As regards synonymy, I have followed my usuai custom,
namely, to give the original reference and a few of the other more
important ones, which, if referred to, will be found to contain a
practically full synonymy.
PRESBYTES NIGRIPES (A. M.-E.).
Semnopithecus nigripes A. M.-Edw. Nouv. Arch. du Mus.
WO Talo JUNIE joe Wy iol WCherAbys lelhyilng di, As So 185 Sschiny, Ess IO:
p- 11 (1875); Anders. Zool. Res. p. 41 (1879).
a. 6 ad. Bali, alt. 250 m., 10th Nov., 1905.
This is an extremely fine example of this scarce species,
agreeing very well with the published descriptions.
PRESBYTES sp. ?
a,b. 2. Nha-trang, 30th Oct., 1905.
Two very young specimens of a species of Presbytes, unfor-
tunately too young for identification.
Nycricesus pyemaus Bonh. (Plate II.)
Abstr. P. Z.S. 1907, p. 2 (Jan. 22, 1907).
Very small, about half the size of . cowcang* Bodd. The
hair is wavy on the body and of a very fine silky texture.
General colour of a uniform orange-rufous, showing no sign of
any dark line down the back or on the head. The under parts,
hands, and feet are lighter in colour and have a silvery-grey
appearance. There is a bare space round the eyes, the muzzle
and lips are white, and a white stripe runs up from the nose
between the eyes to end abruptly on the forehead. The ears are
of moderate size, uniformly rounded, and very sparsely covered
with hairs. The tail is a mere stump.
The skull in its general outline agrees fairly well with that of
NV. c. cinereus from Cochin China, although it is, of course, very
much smaller. In its main characters also it shows no very
* Messrs. Stone and Rehn have pointed out (Proc. Acad. Nat. Sci. Phil. 1902,
p. 138) that the name tardigradus belongs to the Slender Loris “ L. gracilis,” and
that therefore Boddaert’s name must stand for the Slow Loris usually known as
N. tardigradus, ,
1907, | MAMMALS FROM ANNAM. 9)
distinctive points. The molars, however, are conspicuously
different, and enable this species to be easily diagnosed. In
AV. cinereus the first molar is the largest, and the last or third
molar is small and almost quadrilateral in shape. In the present
species, however, the second molar is the largest, while the third
molar is triangular in outline and not very much smaller than the
first molar. In the lower jaw similar conditions obtain, the three
molars are all subequal, the third being slightly the largest,
whereas in iV. cinereus the last molar in the lower jaw is very
markedly smaller than either of the other two.
Dimensions of type from skin (approx.). Head and body
190 mm.; tail 10.
Text-fig. 1. Text-fig. 2.
Text-fig. 1.—Palatal view of skull of Nycticebus pygmeus.
Text-fig. 2.—Lateral view of skull of Nycticebus pygmeus.
Skull. Greatest length 46 mm.; basal length 38; palatal length
17-5; zygomatic breadth 27; interorbital breadth 3; greatest
breadth of brain-case 25; length from palate to lower margin of
foramen magnum 17:5; breadth of basioccipital at its anterior
end 3°7; length of molar series 14.
Hab. Nha-trang, Annam.
Type. B.M. 6.11.6.2. 3. Collected by’ Dr. Vassal on the
13th Nov., 1905.
The small size and peculiar character of the teeth will prevent
this species from being confounded with any others at present
known to exist. Only a single specimen (the type) has been sent,
which is quite adult but not old. It may be noted that in some
respects the teeth tend to approach those in Loris, in which the
second upper molar is larger than the first. In the shape of the
premaxilla also the present species shows a tendency, albeit very
slight, to approach Loris by showing its flat surface laterally
instead of anteriorly. Externally the blaze between the eyes and
its small size are features belonging to Loris, but in the length of
its limbs and general build it is a true Vycticebus.
[=r]
MR. J. L, BONHOTE ON [Jan. 15,
FELIS sp. ?
a. Flat skin with no data of a Cat belonging apparently to the
Felis bengalensis group.
VIVERRA MEGASPILA Blyth.
Viverra megaspila Blyth, J. A. 8. B. xxx. p. 331 (1862).
a. Nambon, Annam.
A fine adult specimen with well-marked and clear-cut spots.
VIVERRICULA MALACCENSIS (Gimel.).
Viverra malaccensis Gmel., Linn. Syst. Nat. i. p. 92 (1788);
Gray, P.Z.S8. 1861, p. 136.
Viverricula malaccensis (Gmel.), Bonhote, Ann. & Mag. N. H.
ser. 7, vol. 1. p. 118 (1898).
a. Imm. No data.
PARADOXURUS MINOR Bonh.
Paradoxurus minor Bonh. Fasci. Mal., Zool. i. p. 9 (19038).
a,b. 9 imm. Bali, Annam, 250 m., 10th Nov., 1905.
These are both very young specimens, which agree closely with
the type.
HERPESTES EXILIs Gerv.
Herpestes exilis Gerv. Zool. de la Bonite, p. 32 (1841); Gray,
P.Z.8. 1864, p. 555.
Herpestes javanicus (Geoft.), Anders. Zool. Res. p. 185 (1879).
Herpestes rutilus Gray, P.Z.8. 1861, p. 136.
Calogale rutilus Gray, P.Z.S. 1864, p. 561.
a,b. 9. Nha-trang, 26th Dec., 1905.
I have carefully compared these specimens with some from
Siam and others from Cochin China, among them Gray’s type
of H. rutilus. The Cochin China and Annam specimens are all
very like each other, and differ in their much redder colour from
Siamese specimens. They also differ in their much deeper colour
from Javan specimens. Gervais’s type of H. ewilis came from
Cochin China, and as his description agrees fairly well with these
fresh specimens, his name of H. exilis, which antedates Gray’s,
must stand.
The skulls of H. emilis, although very similar to those from
Siam and Java, are larger and more robust. The Siamese animal
is apparently intermediate between H. birmanicus and H. ewilis.
The following is a description of the present specimens :—
General colour rufous, punctulated with white. Each hai is
black, with three or four buff or rufous annulations. The distal
annulations and generally the tip of each hair are rufous, while
along the centre of the back, the head, cheeks, and tail these
rufous annulations are deeper in colour and more marked, causing
the animal to appear quite red along those areas. The under
parts are more sparsely covered with hairs and the annulations
1907.] MAMMALS FROM ANNAM. iG
yellowish rather than rufous, except under the chin and at the
root of the tail. The hairs of the tail, more especially under-
neath and at the sides, have long rufous tips.
Dimensions (of Nha-trang specimen, ad. 2 in flesh). Head and
body 364 mm.; tail 284; ear 28.
Skull. Greatest length 78 mm.; basal length 75; zygomatic
breadth 39; palatal length 41; greatest diameter of carnassial 8.
HELIcTIs PIERREI Bonh.
Helictis pierrei Bonhote, Ann. & Mag. Nat. Hist. ser. 7, vol. xii.
p. 592 (1903).
a. Imm. Nha-trang, Annam.
6b. Imm. skull only.
The single skin and skull are too immature to show the dis-
tinctive characters to any marked extent.
TuPAIA Concotor Bonh.
Abstr. P. Z. 8. 1907, p. 2 (Jan. 22, 1907).
Similar in general colouring to Zupaia belangeri. The whole
of the upper parts are of a uniform grizzled greyish-green, each
hair being dark at its base and having one or more buff annulations
and a dark tip. One of the most distinctive features is the absence
of the neck-stripe, so universal among the other species of this
genus. An extremely faint trace of it only is to be made out on
the shoulders, but so faint is it that unless special search be
made it is liable to be overlooked. The tail, which is markedly
distichous, is concolorous with the upper parts, and extremely
thick and bushy. The under parts are somewhat sparsely clothed
with hair; the chin, throat, and breast are uniformly yellow,
while on the other portions the hairs are annulated as on the
upper parts. The bases of the hairs on the under side of the tail
are light.
Skull. In its general character resembles that of 7’. belangeri; it
is, however, slightly larger and witha longer and narrower snout,
in other respects it does not show any marked features.
Dimensions of type (from skin). Head and body 220 mm. ;
tail 140; ear 15; hind foot 45.
Skull. Greatest length 54 mm.; basal length 47; zygomatic
breadth 29; palatal length 27; breadth of skull immediately
behind postorbital processes 17.
Type. B.M. 6.11.6.3. o ad. Collected by Dr. Vassal, 22nd
March, 1906.
Hab. Annam.
Although very closely allied to Tupaia belangeri this species
may easily be distinguished by its larger size, much thicker tail,
and the absence of the light neck-stripe. Two specimens agreeing
in allrespects were brought back by Dr. Vassal. Tupaia chinensis,
described by Dr. Anderson from Yunnan and which is found in
Siam, is rather smaller than 7’. belangeri and consequently quite
distinct from the present form.
8 MR. J. L. BONHOTE ON [Jan. 15,
DENDROGALE FRENATA Gray.
Tupaia frenata Gray, Ann. & Mag. Nat. Hist. ser. 3, vol. vi.
p. 217 (1860).
Dendrogalefrenata Anders. Zool. Res. p.110, pl. 7. figs. 20, 21(1879).
a, 0.
Two very typical examples.
CYNOPTERUS SPHINX (Vahl).
Vespertilio sphina Vahl, Skrivter af Naturhistorie-Selskabet,
Ate Band, lste Heft, p. 123 (1797).
Cynopterus sphinw (Vahi), Bonh. P. Z. 8. 1902, vol. i. p. 38;
id. Fasci. Mal., Zool. pt. i. p. 14 (1903).
a. 6. Nha-trang, Annam, 13th Nov., 1905.
ScoroPpHiLus KUHLII Leach.
Scotophilus kuhli Leach, Trans. Linn. Soc. xiii. p. 71 (1822).
a,b. $. Nha-trang, 10th Oct., 1905.
PETAURISTA YUNNANENSIS (Anders.). j
Pteromys yunnanensis Anders. Zool. Res. p. 282, pl. xxi. (1879).
a. 6 ad. Bali, Annam, 150 m., 10th Nov., 1905.
This individual agrees very well with Dr. Anderson’s description
and plate (quoted above), and I have no alternative but to place
it under his name. At the same time it should be noted that the
typical locality of P. ywananensis is considerably to the north and
that another form of this same species, ?. lylei mihi, is found
in Siam. Jt would therefore appear as if the present race was
in reality a Chinese form and that Annam and Yunnan form its
western limit. Except for the parachute the hairs of the whole
of the back in this individual are tipped with white, but not
sufficiently so as to conceal the chestnut ground-colour.
P. lylei is much darker in general coloration than this species
and the anterior portion of the outer side of the ear is a bright
and pure chestnut.
P. yunnanensis and P. lylei belong to a large group, of which
there are many geographical forms. Until, however, the group is
worked out as a whole, it is best to retain them under binomial
names, but it should be borne in mind that they are merely
geographical forms of a large and widely distributed species.
ScIURUS GRISEIMANUS A. M.-E.
Sciurus griseemanus A. M.-Edw. Rev. Zool. 1867, p. 195;
Bonh, Ann. & Mag. Nat. Hist. ser. 7, vol. vil. p. 274 (1901).
a-d.3 3, 9. Nha-trang, Annam, 26th Dec., 1905.
e. Nha-trang, Annam, Nov. 1905.
f-k. 33,2¢. Hoah Khat, Annam, 26th Dec., 1905.
In. 26, 9. Ninh Hoa, 25th Dec., 1905.
o-p. 6. Bali, Annam, 250 m. alt., 26th Dec., 1905.
The present species and S’. lewcopus* of Gray have hitherto been
confounded and considered as one and the same species. The
* Macrowus leucopus Gray, Ann. & Mag. N. H. ser. 3, xx. p. 282 (1867).
1907. | MAMMALS FROM ANNAM. 9
present series, however, shows that they are really quite good and
distinct species. The most obvious difference is in the colour of
the under parts. In S. griseémanus they are deep chestnut and the
line of demarcation between the upper and under parts is sharply
divided. In S. leweopus, on the other hand, the colour of the
under parts is of a pale rufous buff, which shades gradually into
the grizzled grey of the back. M. Milne-Edwards in his original
description of S. griseimanus distinctly states that the colour
of the under parts is deep chestnut, though females and young
males are sometimes considerably lighter. This enables us to
fix M. Milne-Edwards’s name on the chestnut-bellied form with-
out hesitation. In the present series the colour of the under
parts is very deep chestnut and shows but little variation ; the
two examples from Bali, at an altitude of 250 metres, are, how-
ever, much lighter below, and it may be that these lighter
individuals represent a mountain race of S. griseimanus, but our
material is at present too scanty to settle that question.
S. leucopus differs still further from S. grisetmanus in the
annulations on the hairs of the back being yellower and not of
such a clear grey, thus giving the animal a darker appearance.
The colour of the under parts also extends over the outer sides of
the limbs and is especially noticeable on the thighs.
SCIURUS LEUCOPUS FUMIGATUS* Bonh.
Abstr. P.Z.S. 1907, p. 2 (Jan. 22, 1907).
General colour above similar to that of S. lewcopus, but darker
(see preceding species). Hach hair is very dark, with three or four
yellowish annulations. The tail, which is indistinctly barred, is
similar in colour to the body, but “4he amnulations are of a slightly
deeper tint. Hands and feet dirty yellowish white grizzled with
darker. Under parts and inner sides of the limbs “pale reddish
buff, with the exception of the chin and throat which are grizzled
like the back.
The skulls available are so imperfect that a description 1s not
possible.
Dimensions of type (from skin). Head and body 190 mum. ;
tail 175; ear 17; hind foot 52.
Hab. Ninh Hoa, Annam.
Type. B.M. 6.11.6.25. Collected on the 10th Nov., 1905, by
Dr. Vassal.
This species is easily distinguished from S. lewcopus typicus by
its darker colour above, grizzled hands and feet, and by the outer
sides of the limbs being similar in colour to the rest of the upper
parts. Whereas the typical .S. lewcopus is greyer, the outer sides
of the limbs are buff, and the hands and feet pure yellowish white.
ScIURUS MACGLELLANDI MARITIMUS Bonh.
Sciurus macelellandi maritimus Bonh. Ann. & Mag. Nat. Hist.
ser. 7, vol. iv. p. 51 (1899).
* Since the reading of this paper it has been pointed out to me that the name
“< fumigatus ” 1s preoccupied, haying been used by Gray in 1867. I therefore propose
to rename this squirrel Sciwrus vassali.
10 MR. J. L. BONHOTE ON [Jan. 15,
a. &. One specimen, 10th Nov., 1905.
A single example of this Chinese race has been brought back ;
it resembles the type closely and in all respects. The occurrence
of this form in Annam is certainly surprising; it may, however,
pee aoe ,
prove to range along the whole S. Chinese coast, while S. rodolphi
inhabits the higher ground; on the other hand it might have been
brought over on a ship and escaped.
SCIURUS MACCLELLANDI RODOLPHI A. M.-H.
Seiurus rodolphi A. Milne-Kdwards, Rev. et Mag. de Zool. xix.
p. 227 (1867); id. Rech. Mamm. p. 162 (1871).
Sciurus macclellandi rodolphi A. M.-E., Bonh. Ann. & Mag.
Nat. Hist. ser. 7, vol. iv. p. 54 (1899).
a-c. Three specimens.
These are very typical specimens. The light stripes are of the
same width throughout, and have a tendency to a deeper and more
rufous tinge on their anterior portion. The median dark stripe
tends to become divided down the centre by a brownish grizzled
stripe; the length and extent of this latter stripe seem to be very
variable.
FUNAMBULUS BERDMOREI (Blyth).
Sciurus berdmorei Blyth, J. A.S. B. xvii. p. 63 (1849); Anders.
Zool. Res. p. 261 (1879).
Funambulus berdmoret (Blyth), Bonh, P. Z. 8. 1901, vol. i. p. 56.
a, b. Bali, Annam, 10th Nov., 1905.
S. mouhoti Gray was merely described on a seasonal form of
this species, as I have already pointed out.
FUNAMBULUS RUFIGENIS FUScUS Bonh.
Abstr. P. Z. 8. 1907, p. 2 (Jan. 22, 1907).
Similar to /. rufigenis typicus, but the general tone of colour
very much deeper. Colour above very dark brown, finely grizzled
with buff. The whole of the thighs suffused with deep chestnut,
which is not the case in the typical form. Sides of face chestnut,
rather deeper in tint than in the typical form, and the same may
be said of the chestnut on the under side of the tail. Remainder
of under parts creamy white.
The skull appears to be of a rather stouter build, but without a
good series it would be unwise to lay stress on this fact.
Dimensions of type (from skin). Head and body 190 mm.;
tail (broken) 150; ear 12°5; hind foot 43.
Hab. Bali, Annam, 250 m. alt.
Type. B.M. 6.11.6.28. Collected by Dr. Vassal on the 10th
Nov., 1905.
The much darker general colour and the rufous tinge on the
outer sides of the thighs form good distinctive characters by which
this species may be easily distinguished from the typical race.
Ruizomys pRuiNosus Blyth.
Rhizomys pruinesus Blyth, J. A. 8. B. xx. p. 519 (1851); id.
1907. | MAMMALS FROM ANNAM, 11
Cat. Mamim. As. Soc. Bengal, p. 122 (1863); Anders. Zool. Res.
p. 325 (1879).
a. Plateau of the Lung Brau, 1300 m., 30th Oct., 1905.
This specimen is only a flat skin without a skull; Dr. Vassal
notes that it is “‘not rare” in Annam.
Hysrrix yYUNNANENSIS Anders.
Hystrix yunnanensis Anders. Zool. Res. p. 332 (1879).
a. 2 imm. Ninh Hoa, 25th Dec., 1905.
This is a very young specimen, but has a well- developed nuchal
crest. The skull in its general proportions agrees with Dr.
Anderson’s description. The external characters, however, agree
well with Swinhoe’s /7. swheristata, and, in fact, the only difference
between these two species is to be found in the skulls. In his
original description Swinhoe states that the skull of . suberistata
is indistinguishable from that of MH. hodgsoni Gray, which, of
course, is quite distinct from Anderson’s species. Unfortunately
there are no specimens from China in the British Museum which
would enable us to determine definitely whether there be two
Crested Porcupines in China, or whether Anderson’s and Swinhoe’s
species are in reality one and the same.
Lepus vAssati Thos.
Lepus vassali Thos. Ann. & Mag. Nat. Hist. ser. 7, vol. xvi.
p. 425 (1906).
9. Nha-trang, Annam, 25th Dec., 1905.
Mr. Thomas having recently described this specimen, I need
only refer those interested to the paper quoted above.
TRAGULUS KANCHIL AFFINIS Gray.
Tragulus affinis Gray, P.Z.S. 1861, p. 138.
Tragulus kanchil pierret Bouh. Ann. & Mag. Nat. Hist. ser. 7,
vol. xi. p. 293 (1903, 1st March).
a-c. 3. Nha-trang, 22nd March, 1906.
aie Specimens agree in all respects with my description of
. k. pierret quoted “above. I have used Gray’s name for this
Hess in preference to my own, as Mr. Miller has pointed out to
me that Gray’s 7’. affinis was chiefly based on specimens from
Cochin China; and Mr. Miller having, previously to my paper,
described the Peninsula form under the name 7’. ravus, Gray’s
T. affinis became ipso facto restricted to the race from Cochin
China: with this finding I quite agree.
MANIS JAVANICA.
Manis javanica Desm. Mamm. p. 377 (1820); Anders. Zool.
Res. p. 352 (1879).
a. Dang-trang, near Nha-trang, 25th Dec., 1905.
EXPLANATION OF PLATE II.
Nycticebus pygmeus, p. 4.
12 MR. P. H. BAHR ON THE (Jan. 15,
2. On the * Bleating” or “ Drumming” of the Snipe
8 g P
(Gallinago celestis). By P. H. Baur, B.A., F.Z.8.
[Received November 20, 1906. |
(Text-figures 3-9.)
This subject has been much discussed, but the interest taken in it
seems rather to have waned during the latter years. I think there-
fore it would be profitable to inquire once more into this strange
phenomenon, especially as many points require elucidation, in the
explanation of which authorities differ considerably. I believe it
is well known to all of us that during the breeding-season our
Common Snipe performs certain aerial evolutions, producing at the
same time a mysterious sound, called in various parts of the country
bleating, humming, drumming, or whirring, ‘“‘ Meckern” in Ger-
many, while in parts of Scotland the popular name of the bird is
‘* Heather Bleater,” and in France “ Chévre volant.” The process
is Shortly as follows :—The bird is seen to fly straight up to a
height of from 60-100 feet, then, turning, to spread its tail, close
its wings, and drop to within 20-30 feet of the ground, producing
at the same time this mysterious sound, which has puzzled observers
so. As to the cause of it many theories have been put forward
by scientific ornithologists, sportsmen, and foresters. We may
group the evidence for these theories under four heads :—
I. The sound is produced by the vocal organs.
II. The sound is produced by the rectrices of the tail, which I
hope to be able to prove is correct.
Il. The sound is produced by the action of the primaries of the
wing.
IV. The sound is produced by the combined action of the wings
and tail (maintained by a few observers).
I. The first evidence in favour of this theory which I can find
is an article by Débel (‘Jiiger Practica,’ 1783, pt. i. p. 73), who
says that the Snipe produces at night-time, while sitting on the
ground in a marsh or close to water, a noise which the ignorant
would mistake for the bleating of a young goat.
Bechstein, in 1789 (Naturgesch. Deutsch. 2nd ed. vol. iv. p. 190),
maintains “that the Snipe makes its plaintive ery, like a goat
bleating, with its beak and not, as has been lately affirmed, with
its wings.” He adduces evidence of birds bleating whilst perched
on the tops of trees.
Hintz (‘ Naumannia,’ 1854, p. 290), from observations made
from 1816-19, believes the sound is made by the bill. He heard
Snipe “bleating” whilst sitting on top of withered oak-trees, first
uttering their call-note “ pecka pecka,” and then bleating.
Zoppritz (Ornith. Centralblatt, Nov. 1880) seems to be very
certain of the accuracy of his observation, for he writes :—‘ Two
years ago I published an article on this subject in a sporting
journal, wherein I offered to pay a fine of 500 marks to the treasury
of the Allgemeine Deutsche Jagd-schutz-verein if three umpires
appointed by the Verein publicly declared that they were convinced
1907. ] ‘‘BLEATING” OF THE SNIPE. 13
that Snipe produced their bleating notes, not through the vocal
organs, but by means of their wings, with or without help of their
tail-feathers.” Nobody seems to have accepted the offer, and we
understand Herr Zéppritz kept his 500 marks. Again, we find
in the same paper :—‘ Anyone with a knowledge of mechanics or
physics, if he sufficiently examine a dead Snipe, must be convinced
that so small a bird, with wing- and tail-feathers so comparatively
weak, cannot possibly produce sounds with them, which are at
such a distance so sharply accentuated. Hence [indulge the hope
that the adherents of the wing and tail theories will now with-
draw their opinions and acknowledge that to err is human.”
Lastly, in Seebohm’s ‘ British Birds,’ vol. ii. p. 244 (1885), we
find the following:—‘‘T have listened to the drumming of the
Snipe scores of times with the express purpose of discovering the
mode in which the sound is produced, but must confess myself
completely puzzled. Arguing from analogy, I should say it was
produced by the vocal organs, and is analogous to the trill of the
Stints and other Sandpipers. The fact that it appears to begin
the instant the bird begins to descend induces me to think that,
after allowance is made for the time it takes for sound to travel,
it must really begin before the descent, whilst the bird is not
moving very rapidly.”
Of such is the evidence with which we have to deal. At
the present day I trust this theory has but few adherents. Pralle
seems to have disposed of it entirely by a note recorded in ‘ Nau-
mannia’ (1852, pt. 1. p. 25), that on 24th March, 1846, he heard the
Snipe utter its note, “ gick-jack, gick-jack,” while bleating.
If. That such a sound could be produced by such feeble instru-
ments as the rectrices seems to have been foreshadowed by
Naumann, curiously enough by a misprint of the word “tail” for
‘“‘ wing-feathers ” (‘ Federwild-jagd,’ von Louis Liegler, Hannover,
1846, p. 174) :—“ It is not hard with sharp eyes (still more with
field-glasses) to observe the quivering motion of the tips of the
tail-feathers [the italics are my own] during each downward and
upward flight through the air, sufficient to convince one that the
sound is thus produced, and not from the throat of the bird. ‘The
sound, or at least a similar one, can be produced if one take the
primaries of certain (but not too small) birds and fasten them to
the end of a long cane, and strikes with this, as with a sword, ina
draught of air.”
To which Jickel (‘ Naumannia,’ 1855, pp. 112, 113) replies and
casts doubt on Naumann’s theory (or, rather, mistake) of the sound
being produced by tail-feathers.
In 1858 Mr. Wolley communicated a paper by Mr. Meves, of
Stockholm (Proc. Zool. Soc. Lond., April 1858, p. 199), wherein
Mr. Meves, in consequence of the misprint already quoted, was
led in 1856 to make experiments with the rectrices of G. celestis.
He remarked with surprise “that the humming sound could
never be produced whilst the bird was flying upwards, at which time
the tail is closed; but only when it was casting itself downwards
in a slanting direction, with the tail strongly spread out.”
14 MR. P. H. BAHR ON THE (Jan. 15,
He then examined the tail-feathers of our common species
more closely and found “the jist (outer feather) especially very
peculiarly constructed ; the shaft uncommonly stiff, sabre-shaped.
The rays of the web stro a bound together and very long, the
longest reaching nearly ? of the whole ‘length of the web,
like the str ings of a musical instrument. If one blows from
the outer side upon the broad web it comes into vibration, and a
sound is heard, which, though fainter, resembles very closely the
well-known neigh.” He then fastened the outer or first feather
with fine thread to a piece of steel wire and fixed it to a 4-foot
stick, and found if he drew this with the outer edge of the feather
through the air, at the same time making shaking motions of
the arm to represent the shivering of the wings during flight,
he was able to produce the neighirg sound with astonishing
exactness. In bringing the matter before the Zoological Society
Mr. Wolley confirmed Meves’s experiments. These experiments I
hope to explain more fully anon.
John Hancock, in 1875, in the ‘ Birds of Northumberland and
Durham,’ vol. vi. pp. 105-113, severely criticises Meves’s theory
and experiments.
I cannot quote his article at length, for it is a very long one.
He argues, from the diversity of structure exhibited by the rectrices
of various species of Snipe, that they cannot be musical instruments.
He failed to produce the neighing sound of a Snipe by Meves’s
experiment, but admits “‘ when the web of almost any firm feather
is blown upon a low vibrating sound is produced ; and such a sound
is stronger when a tail-feather of the Common Snipe is used,
arising apparently from the fact that the inner web is wide and firm ;
but the sound is so low that it cannot be heard many yards off.”
Later: ‘The sound is audible at a great distance, even when
the bird has risen high into the air. No sound that could be
produced under any circumstances by such feeble instruments as
the lateral tail-feathers of the Snipe, instruments not larger than
the wings of a Dragonfly, could be heard at any considerable
distance. And it is scarcely to be doubted by anyone that the
wings of a Snipe vibrating rapidly will produce some sound louder
than any that could be made by a pair of small tail-feathers of a
bird rushing downward through the air.”
Prof. Altum (‘Ornithologisches Centralblatt,’ Oct. 1880) satisfied
himself that he could produce the sound with the lateral tail-
feathers. He, however, quotes “two adverse cases ” :—
(1) A Snipe was observed “ bleating” as it sat, or rather stood,
on an elevation.
(2) A certain Alex Schmidt winged a Snipe which, with tail
stiffly expanded, began to bleat in his hand, the air blowing
through the web of the feathers, the wings being held
close to the bird’s side. Every time the bird was moved
rapidly against the wind his object was attained.
Tt is to be noted that in both cases a strong wind was noticed
to be blowing. ‘That these two cases are easily explicable I hope
to adduce evidence later on.
1907. | ‘‘BLEATING” OF THE SNIPE. 15
In answer to this paper, von Zoppritz (Ornith. Centralblatt,
Nov. 1880) disputes Altum’s tail-theory, giving four reasons,
which JI will not here quote*.
III. The wing-theory has had the greatest number of adherents.
Macgillivray in 1840 (‘ British Birds,’ vol. iv. p. 372) expresses
his conviction that this is the case. Sir Wm. Jardine, in the
‘Naturalist’s Library’ (vol. xxvi. Ornithology, p. 180), says: ‘‘The
sound is never heard, except m the downward flight, and when
the wings are in rapid and quivering motion. ‘Their resistance
to the air without doubt causes the noise. Dr. Saxby (‘ Birds of
Shetland, p. 204) expresses his conviction that ‘ drumming is
produced by the vibrations of the wings alone.’ ”
Naumann, in the article already quoted, and Zoppritz (Ornith.
Centralblatt, Nov. 1880) wrote in favour of it in Germany.
John Hancock, whom I have already quoted, agrees in the
main with what Mr. J. E. Harting (‘ Essays on Sport and Natural
History,’ pp. 284 ef seq.) has written :—‘‘ From the peculiar
vibration of the wings in the downward descent of the bird, it
would appear that the primaries, instead of firmly overlapping
each other, are, in the act of humming, turned broadside in the
air, which is thus able to play across the inner web of each, and so
to impart to each a vibratory motion and consequent sound—faint
indeed in the case of a single feather, but audible enough when
an entire wing is acted upon. Whether this be the true expla-
nation of the singular sound, it is, of course, not easy to prove
conclusively ; but it has certainly been accepted as such by many
naturalists in England, who are the more inclined to adopt this
view from having observed Peewits, Rooks, Gulls, and other birds,
with tails very different from that of a Snipe, make an analogous
sound while falling through the air.” That Mr. Harting was
partially successful in producing the bleat artificially is evident, for
he has ‘“‘ succeeded beyond expectation in producing a sound like
the ‘ humming’ of the Snipe.” Again: “ But any of the primary
wing-feathers will give forth a faint sound, which may be increased
in proportion to the number of them passed through the air at
once.” But he finds that the tail-feathers when fastened into a
switch do not occupy the position they do naturally in the bird’s
tail, because they are drawn through the air at right angles to the
direction of flight, ‘‘in a position which is occupied naturally by
the primaries, but unnaturally by the tail, and hence it must be
the primaries (collectively) which produce the sound in nature.
In this our sense of hearing is assisted by the sense of sight, for
a perceptible vibration of the quill-feathers is observed every time
the bird descends.”
IV. Two observers maintain that the sound is produced by
the agency of both the wing- and tail-feathers.
H. Gadumer (‘ Naumannia,’ 1853, pp. 411-413) watched a bird
* (Mr. F. W. Headley (° Nature’, vol. Ixx. p. 103, 1904) supported the theory that
the drumming was produced by the outermost tail-feathers, and adduced an expe-
riment by which the sound could be produced artificially. |—Ep. P. Z.S.
16 MR. P. H. BAHR ON THE (Jan. 15,
bleating, with a good field-glass. “The air pressing between the
wings and the tail (a natural parachute) causes all the feathers of
the tail and wings to vibrate and gives rise to the bleating sound
—which is modified by stronger or weaker vibrations.”
Text-fig. 5.
a. Snipe bleating, showing characteristic position.
b. Formation of tail as ordinarily held in flight.
1907. ] ‘““BLEATING” OF THE SNIPE. 17
Capt. W. V. Legge, in his appendix to the ‘ Birds of Ceylon,’
expresses the opinion that the sound is produced by the combined
action of the wings and tail.
I have dwelt on the literature for some length, in order that
one may review the evidence adduced by the adherents of the
different theories.
In the summer of 1904, in the Fens of Cambridgeshire, I began
to observe the Snipe in the act of bleating through a strong prism
binocular, I had read none of the literature on the subject, and
so had no preconceived ideas. The observations I made then I
have had ample opportunities of confirming.
I find that ordinarily the bird flies up to a height of 60-100 feet
above ground, in windy weather going higher, with its tail held
in the ordinary position of flight (text-fig. 3, 6), then, turning, it
spreads its tail out like a fan, the two outer tail-feathers being
spread out well in front of the other twelve and held firmly there
(text-fig. 3,a). Immediately the bird begins to descend the bleat
is heard (making due allowance for the time it takes for sound to
travel). While descending the bird makes tremulous motions
with its wings from the radio-carpal joint. The descent is made
from 30-40 feet and occupies 2-3 secs., the bleat lasting the same
time. The bird does not drop head foremost through space, but
at an angle of from 45°-60° with the horizon. The tail as a whole
is not vibrated, but it is quite easy to see the two outer tail-feathers
with a strong glass vibrating to such an extent that their terminal
portions become indistinguishable. Snipe begin to bleat in March,
but if the weather is mild, in February, and continue to the end
of May, though I heard one last year in Sutherland still bleating
on June 25th.
At the beginning of the breeding-season they may be seen
bleating in pairs; but later on, when the hen is sitting, the cock
bird may be seen performing alone over the marsh where the nest
is placed. Under favourable conditions many bleat together,
circling round the same spot for hours. On April 12th of last
year, I had the good fortune to hear no less than twelve birds
bleating together, a concert which they kept up all through the
night. Every now and again, as if by common consent, there
would be a lull, and all the birds would settle, but directly one
began again ali the rest immediately joined in the chorus.
Snipe bleat best in the early morning and in the evening,
especially when the weather is dull and damp. It may be of
interest to note that last spring I saw a specimen of the melanistic
variety (Sabine’s Snipe) bleating.
Once having convinced myself that the two outer tail-feathers
are invariably spread out beyond the others, a fact which is now
obvious to me with the unaided eye, it seemed to me that the two
outer tail-feathers must be the active agents in causing the bleat.
I accordingly procured several tails of the Common Snipe, and
taking the two outer tail-feathers, pierced the shaft with a pin,
Proc. Zoou. Soc.—1907, No. II, 2,
18 MR. P. H. BAHR ON THE [Jan. 15,
to which I firmly bound it with cotton and inserted the feathers
into a cork at the end of a stick some six inches long. A hole is
bored at the other end of the stick and a long string attached.
This is whirled round the observer’s head and a typical bleat is
produced. The second outer tail-feathers (sixth pair) produce a
fainter sound, though this varies much in individual tails, the
others make no sound at all.
Text-fig. 4,
Varieties of outer tail-fegthers of Gallinago celestis. (Natural size.)
1907.] ‘““BLEATING” OF THE SNIPE. 19
In order to ensure the success of the experiment it is necessary
(1) that the feathers be placed so that the narrow edge, the outer
web, shall encounter the resistance of the air; (2) that the feather
be firmly bound to the pin, so that it cannot turn on its support ;
(3) that the string be tied to one end of the stick, so that the
long axis of the stick makes an angle with the direction of the
string, if I may so put it, so that a vibratory motion is imparted
to the stick as a whole, thus simulating the tremulous motion
of the Snipe’s wings during the descent; (4) lastly, that the
apparatus be moved at a uniform rate and not too fast.
It is then found that after a period of silence the feathers begin
to vibrate : first, the long-drawn-out note, which I may represent
as “whti whtutu,” becomes gradually audible, it is then succeeded
by a series of high and low notes ‘ bah-bah-ah-ah,” resembling
the bleat of a young goat, lasting 3-5 secs., followed by a pause
of equal length. This is repeated as long as the apparatus is
revolving at a uniform rate. It is found that the individual tail-
feathers, of which I collected a good number during the winter,
vary considerably both in size, breadth, and markings, and, as
might be expected, the note produced varies according to their
physical characteristics. Thus a long narrow feather produces a
sound of far higher pitch than a broader one of the same length
(vide text-fig. 4). This fact I have noted when comparing the
sound made by several birds when performing the nuptial evo-
lutions over their breeding-grounds. To ascertain which part of
the feather is essential in the production of the sound, I have cut
off the narrow outer web, without altering the bleat in any way ;
but if the barbs of the inner web be so disarranged that there is
a break in their continuity, the web ceases to vibrate, and no
sound is produced. That the vibration of the inner web is the
active causative agent may be seen by the following simple
experiments :—The feathers are attached to a cork, with the outer
web held away from the observer, so that the narrow outer web
shall cleave the resistance of the air. Thus affixed they are held
out of the window of a train, or while riding a bicycle. As the
resistance of the air is encountered the inner web begins to
vibrate, slowly at first, but as the train gains speed, so rapidly
that its outline is entirely lost, and it becomes a blurr; a low
humming sound is at first heard, which soon reaches the typical
pitch of the bleat. When the train has reached the speed of
some 20 miles an hour, the whole feather will vibrate on the pin.
If the feathers are at all loose on their pins it 1s curious to observe
how they will always turn round so that the narrow outer edge
encounters the resistance of the air. Furthermore, if the feathers
be damped, they appear to act better, thus explaining, perhaps,
why Snipe are found to be lable to bleat in damp weather. I
-think this simple experiment readily explains away the ‘“ adverse
eases ” of Prof. Altum (‘ Ornithologisches Centralblatt,’ Oct. 1880)
already mentioned.
That the bens bleat as well as the cocks is now, I suppose, a
well-known fact (cf. von Preen, ‘ Naumannia,’ 1856, pp. 426, 427,
O*¥
rai
20 MR. P. H. BAHR ON THE [Jan. 15,
and Meveg, Proc. Zool. Soc. 1858, p. 200). I have observed it on
several occasions myself. In the summer of 1902 I found four
newly hatched Snipe in a patch inhabited by only a single pair;
while lying concealed in the neighbourhood I observed repeatedly
both old birds drumming above me. From the similarity of
Text-fig. 5.
a. Dorsal view of musculature of tail of Snipe.
C=levator coccygis.
- D=pygostyle.
E=fused spines of caudal vertebre.
A=slip of m. ilio-coccygeus to outer
rectrix.
B=m. ilio-coccygeus.
b. Ventral view of musculature of tail of Snipe.
D=depressor coccygis.
A=m. pubococcygeus ext.
K=pygostyle.
B=m. pubococcygeus.
C=m. caud. ilio-femoralis.
structure of the tail-feathers in both sexes, a fact which I have
ascertained by dissection, one would infer that both sexes
drummed. I cannot, however, agree with Meves that “as the
feathers of the hen are generally less than those of the cock bird,
the noise also made by them is not so deep as in the other case ”
1907. | “‘ BLEATING” OF THE SNIPE. 21
(op. cit. p. 200). I can find no difference either in the length of
the feathers or in the intensity of the sound produced by the
feathers of either sex. I have received a letter from Mr. 8. A.
Buturlin, in which he says that in 1905, on the Kolyma Delta, he
frequently observed both sexes of the eastern representative of
our species (Gallinago raddit) drumming.
Since the two outer feathers are extended beyond the other
twelve during the descent, as I have described, I sought to find
by dissection a mechanism by which this might be produced. On
examining the tail of a freshly-killed bird, it is quite easy, by
spreading out the tail, to make it assume the arrangement shown
(text-fig. 3). I was unable, however, to find any special muscle
peculiar to the species controlling the outer two tail-feathers.
The muscle pubococcygeus ext. (text-fig. 5, 6) is inserted into the
base of the shaft of the outer two tail-feathers, and is quite capable
of performing this function. This muscle is to be found equally
well developed in the other species of Plovers and Waders which
IT examined. The nomenclature of the muscular system of the
tail is that of Gadow in Bronn’s ‘ Thier- Reich.’
I have tried the same experiments as I have just described with
the primaries from the wing of the Snipe, and was not able to
produce any more sound with them than with others taken from
other kinds of Waders, Pigeons, &e. There seems to have existed
an opinion at one time that the bird produces two sounds, one
with the wings and the other with the tail, the former being
known as humming or drumming, and the latter whirring or
bleating, produced while the bird is on the ground (cf. ‘ Zoologist,’
1881, p. 212, and 1846, p. 1501). I cannot say that this agrees
with my own experiences.
An Hxamination of the Structure of the Tail of Gallinago ccelestis
(text-figs. 6, A, and 7).
The normal number of feathers in the tail of this species is 14.
It could hardly fail to strike the observer, on examining the tail,
that the outer two differ considerably from the rest. Firstly,
they are lighter in colour and their texture is firmer. On closer
examination the shaft is seen to be strong and firm, presenting a
decided outward curve towards its lower third. The outer web
is narrow, and formed of stiff rami, which can easily be separated.
The inner web, on the other hand, is extremely broad, being six
times as broad as the outer, and formed of long stiff rami, of which
some reach quite three-fourths the whole length of the feather,
making a very acute angle at their insertion with the stem (text-
fig. 6). The individual rami adhere firmly to one another, and
can with difficulty be separated. These are provided with two
well-developed rows of radii, the distal and the proximal rows
(text-fig. 7, B), the former are twice the length of the latter. I
must here express my great indebtedness to Mr. W. P. Pycraft,
who has allowed me to make full use of his excellent paper, on
Zo MR. P. H. BAHR ON THE [Jan. 15,
“The Interlocking of the Barbs of Feathers” (‘ Natural Science,’
vol. ii, No. 19, Sept. 1893), and from which the illustration (text-
fig. 7, A) is copied. Under the microscope the distal row is seen
Text-fig. 6.
A. Half of tail of Gallinago celestis from without inwards left to right.
B. Half of tail of G. delicata from without inwards left to right.
C. Half of tail of G. gallinula.
to be well provided with hamuli and cilia (text-fig. 7, B). The
hamuli deserve attention (text-fig. 7, C), since I believe them to
1907. ] ‘“BLEATING” OF THE SNIPE. 23
Text-fig. 7.
> 3 3
> ; \ ll
MIE Wii) Y , ij py yp he Y) gr
H]
WI
x
Wr hamuli
|
A. Section of two rami of a feather showing interlocking of distal and proximal
radii. (After W. P. Pycraft.)
B. Ramus of Gallinago celestis, showing proximal and distal rows of radii.
C. Distal radius of G. celestis. D. Proximal radius.
K. Distal radius of middle tail-feathers of G. celestis.
24. MR. P. H, BAHR ON THE [Jan. 15,
be the essential factor in producing the bleat, in that they hold
the stiff rami together like the strings of a harp. They are
seven or eight in number, a number in excess of any other
species of Snipe, and are well-formed, possessing a well-hooked
terminal portion, which interlocks with the upturned edge of the
radi of the proximal row (text-fig. 7, A & D). The outer web is
formed of stiff rami, which possess rudimentary radii unprovided
with hooklets.
Of the remaining feathers, the sixth pair most nearly approaches
our type feather in structure (text-fig. 6, A). The shaft is, however,
not so strong, the outer web is broader, the inner narrower, and
the rami are not so long, nor do they form such an acute angle
with the stem (text-fig. 6,A). The hamuli are fivein number and
not so well-formed, and, as I have said before, the sound produced
by the vibration of the inner web of these feathers cannot compare
in intensity with that produced by the outer pair. Thus the
outer web becomes gradually broader, the inner gradually
narrower as we reach the central tail-feathers (text-fig. 6, A), and
the rami become progressively weaker, the hamuli fewer in
number. Thus the distal radius of the middle tail-feathers
possesses but four feebly curved hamuli (text-fig. 7, E).
I have examined the tail of this species during the moult. On
the 17th of August, 1906, I received several from Scotland, just
at the beginning of the moult. The outer tail-feathers, I find,
have lost much of their bleating power and the note produced is
not so intense. On microscopical examination I find the cilia
(text-fig. 7, C) have all been worn away. From this I infer that
the cilia play a certain part in the production of the sound.
From the 17th August to the 6th September I received
many tails in which the new feathers were just growing, and I
find that in every case the outer tail-feather (the sonorous instru-
ment) is the last to be assumed. The newly assumed feather
possesses fu]l bleating powers. I have also examined feathers
from young birds of the year directly they have assumed their
full plumage; these, I find, will bleat as well as those of a fully
adult bird, and possess the normal structure and characteristic
number of hamuli.
One variety of G. celestis still remains to be mentioned, i. e.
Gallinago raddw (Buturlin), which is the eastern representative
of our species and is much lighter in colour. With characteristic
kindness, Mr. Buturlin has sent me skins of this and several other
species from the Kolyma Delta in Siberia (69° 4’ 20” N. and
160° 55" E.), accompanied by most valuable notes, for which I am
deeply indebted. The feathers of the tail, of which I have figured
a specimen (text-fig. 4, 4), behave in the same way as those of our
species.
Examination of the Tail of other Species of Gallinago.
Gallinago delicata, or Wilson’s Snipe, of N. America, differs
in the eyes of some materially from G. c@lesiis in possessing
Sa) “BLEATING” OF THE SNIPE. 25
sixteen tail-feathers. Of these the outer two are specialised, but
differ from those of G. celestis in that the mner web of these
feathers instead of being broader is narrower than that of the
other (text-fig. 6, B); they are, in fact, somewhat attenuated—a
process which, as we shall see later, reaches its extreme in the
Pin-tailed Snipe (G’. stenwra) of India. The shaft is strong, but
not so strong as that of our species. The inner web is three
times as broad as the outer. Both feathers will produce a
bleat on experiment; the sound is of a far higher pitch than
that of G. calestis, as might be inferred from the character of
the feathers, and is what is aptly described by the Americans
as “winnowing.” The rami are shorter in comparison with
G. celestis, and make an acute angle with the shaft. The radii
of the outer web are rudimentary, of the inner web the distal
is but one-third longer than the proximal row; the hamult
are five in number, but are not so well hooked as those of
G. celestis. Of the two, the outer or eighth is more attenuated
than the seventh pair.
Regarding the habits of this species I have the following
references :—
Baird, Brewer, and Ridgway, ‘ Water-Birds of N. America,
vol. i. p. 191 :—‘“‘ Capt. Blakiston noticed that this species per-
formed the same evolutions as the European bird, this usually
about sunset, but at times continuing 13 hours later. The noise
made on these occasions he compares to rapidly repeated switches
of a cane in the air, and this was repeated every half minute with
cecasional longer intervals. The sound lasted about three seconds,
and was made as the bird descended rapidly in a vertical direction,
being caused apparently by the quill-feathers of the wings. This
sometimes took place in the middle of the day, but only during
the love season.”
Again :—
Audubon, ‘ Birds of America,’ p. 343 :—‘ These birds are often
met with in meadows, or on low grounds, and by being on the
spot before sunrise, you may see both mount high in the air in a
spiral manner, now with continuous beats of the wings, now in
short sailings, until more than a hundred yards high, when they
whirl round each other with extreme velocity and dance as it
were to their own music, for at this juncture, during the space of
5 or 6 minutes, you hear trolling notes mingling together, each
more or less distinct, perhaps according to the state of the
atmosphere. The sounds produced are extremely pleasing, though
they fall faintly on the ear, but I am well assured that they are
not produced simply by the beatings of the wings, as at this time
the wings are not flapped, but are used in sailing swiftly in a
circle not many feet in diameter. A person might cause a sound
somewhat similar by blowing rapidly alternately from one end
to another across a set of small pipes consisting of 2 or 3
‘modulations.”
From this we gather that this species performs its evolutions at
26 MR. P. H. BAHR ON THE [Jan. 15,
a greater elevation than our species, also that Audubon noticed
the fact that both cock and hen bleat.
Wilham Brewster, in Chapman’s ‘ Handbook of Birds of North
America,’ writes, pp. 154-155 :—‘‘In the springtime, and occa-
sionally in autumn also, Wilson’s Snipe mounts to a considerable
height above his favourite meadows, and darts downward with
great velocity, making at each descent a low yet penetrating
tremulous sound, which suggests the winnowing of a domestic
Pigeon’s wings, and if heard at a distance, the bleating of a goat,
and which is thought to be produced by the rushing of the air
through the wings of the Snipe. This performance may be
sometimes witnessed in broad daylight, when the weather is
stormy, but ordinarily it is reserved for the morning or evening
twilight or for moonlight nights, when it is often kept up for
hours In succession.”
Other American species deserve mention here :—
Gallinago nobilis (text-fig. 8, B) has 16 tail-feathers, of which
the outer three are attenuated and the fourth partially so. It
inhabits Ecuador and Colombia, and is allied to G. australis,
which species I shall treat of later. I can find no reference to the
breeding-habits of this species. On experiment the three outer
feathers bleat well. As in G. delicata the rami of the outer web
possess but rudimentary radii; those of the inner, however, possess
a distal row which is one-third longer than the proximal, and the
former is provided with five hamuli which are not well-hooked.
The rami are thicker and stiffer than in the aforementioned species,
and I suspect it is more by the vibration of the rami as a whole
that the sound is produced, in contradistinction to the vibration of
the inner web alone as in @. celestis and delicata. The inner web
of the eighth pair is but little broader than the outer, but that of
the seventh pair is nearly twice as broad, thus resembling the
outer tail-feathers of G. delicata. The middle tail-feathers conform
to the ordinary type. Thesound produced is hard to describe; it is
flute-like, but possesses a definite bleating character.
Gallinago frenata.— Another South American species inhabiting
Brazil, believed to be a Neotropical form of G. delicata. This
species has 16 tail-feathers, of which the outer four are
attenuated, the outermost being one-fifth inch in diameter, the
inner being but slightly wider than the outer web. The inner web
becomes progressively larger towards the centre of the tail. . This
Species agrees with the foregoing in having the specialised
feathers of a lighter colour: all four bleat; this is similar to that
of G. nobilis, but shriller: microscopically they resemble the
structure of G. delicata. The distal radii of the inner web are one-
third longer than the proximal row and possess five well-curved
hamuli. The rami of the outer web are stiff and structureless.
G. paraguay is a subspecies of G. frenata inhabiting Paraguay.
It is much larger than the type, and the tail-feathers are
1907.] ‘‘BLEATING” OF THE SNIPE. 27.
consequently larger and the bleat deeper in tone ; otherwise they
agree in number and structure. A description of the breeding-
Text-fig. 8.
A. Half of tail of @allinago major from without inwards left to right.
B. Half of tail of G. nobilis from without inwards left to right.
C. Half of tail of G. stenwra.
habits of this species is to be found in a paper by Durnford in the
‘This, 1877, p. 198:—‘ During the spring they go through the
28 MR. P. H. BAHR ON THE Jan. 15
?
same aerial movements as the Common Snipe at home, rising to
a great height by a circling motion, and ‘drumming’ whilst
descending in a diagonal line. How is this curious habit to be
accounted for in the South-American and European forms, except
by the theory of inheritance from a common progenitor ?”
Gallinago australis —Latham’s Snipe has 18 tail-feathers, of
which the outer three are attenuated, two being less than 3 of an
inch in diameter; the outer six, however, are doubtlessly specialised,
as they differ markedly from feathers from the centre of the tail
both in structure and colour. The feathers bear a certain resem-
blance to those of the South-American species. The shaft is
thick and curved ; the rami of the inner web are long and thick,
but more easily separated than in the latter species. The rami
are peculiar, in that they are thicker and stiffer than in any other
species. The distal rami of the inner web are longer than
the proximal row and are provided with 5 hamuli. The rami of
the outer web are stiffand structureless, thus resembling G. celestis,
paraguaye, and frenata. The feathers produce aloud bleat, some-
what similar to that of G. celestis.
I have received from Mr, Alan Owston, of Yokohama, a skin of
this species, accompanied by some very valuable notes, which
add materially to our knowledge of the habits of this species, for
which Iam greatly indebted to that gentleman. He says: ‘“ They
breed on the grassy moorland at the foot of Mt. Fugiyama, at an
elevation of 2000-3000 ft. above the sea (Fugiyama is 12,500 ft.
high). I have watched them on the 28th April, and on other
dates during the breeding-season. When alarmed they fly round
overhead, circling round generally against the sun, and every now
and again they begin to cry ‘chip, chip, chip, sheep, cheo, che-
cheo,’ and then rush downwards at the intruder, beating the air
in the descent and making a terrific rushing noise.”
He also sends me an extract from Capt. T. W. Blakiston’s
notes on the breeding-habits of this species published in the
‘Chrysanthemum’ for Nov. 1882, p. 524 et seg., referring to
“Birds observed on the 8.E. coast of Yezo in May” :—“ The
Australian species act very like the Snipe of North America,
by flymg round pretty high and making sudden rapid descents
almost to the ground, which latter movement is accompanied by a
whisping noise. At evening and during the day in dull weather,
these evolutions are commonly performed; and in dirty rainy
weather the noise is heard even in the middle of the night.”
Gallinago aucklandica.—Resembles G. australis in having 18
tail-feathers, the outer three of which are attenuated; they are,
however, much softer in structure than in that species. The rami
of the inner web are easily separated, and possess 4 hamuli; those
of the outer are provided with rudimentary rows of radii, thus
approximating to certain Asiatic members of the genus. I have
received a specimen of this rare species from the Christchurch
Museum, N.Z. No mention of any bleating-habits is made in
1907.] ‘“‘BLEATING” OF THE SNIPE. 29
the literature. Contrary to what one would expect from its
peculiar Rail-like appearance, the tail-feathers produce a very
distinct and pleasing sound of a high-pitched character, some-
what resembling that of certain Asiatic species.
Gallinago equatorialis (nigripennis) is an inhabitant of Central
Africa, south and east of the great desert. It possesses 14 tail-
feathers, of which the outer four are attenuated, and are less than
1 inch in diameter ; they are pure white. This form is nearly allied
to the Common Snipe. I have been quite unable to procure either
any feathers or a skin of this species.
Its habits are described in Reichenow’s ‘ Végel Afrikas,’ Band i.
p- 236 :—“ This bird is called Spook Vogel by the Boers, on account
of its drumming cry, which the bird makes in the morning and
evening during its flight.”
Gallinago gallinula.—The Jack Snipe has 12 tail-feathers, of
which the outer three are markedly shorter than the three central
ones (text-fig. 6, C).
Their texture is soft and the rami are easily separated, in
contradistinction to those of the species we have already con-
sidered. On experiment these feathers produced no sound at all.
The structure of the outer web of the outer feathers more
nearly approaches that of the inner—a marked difference to that
found in the other feathers we have been considering ; that is, the
rami of the outer web are provided with distal and proximal rows
of radii and thus adhere together. The distal radii are provided
with 4 hamuli both in the outer and inner webs.
The breeding-habits of this species were described originally by
Wolley in Hewitson’s ‘Eggs of British Birds,’ vol. ii. p. 356 :—
“Tt was on the 17th June, 1853, in the great marsh of Muonio-
niska (Finland), that I first heard the Jack Snipe, though at that
time I could not guess what it was—an extraordinary sound, unlike
anything I had heard before ; I could not tell from which direction
it came, and it filled me with a curious surprise. My Finnish
interpreter thought it was a Capercally, and at that time I could
not contradict him. I know not better to describe the noise than
by likening it to the cantering of a horse in the distance, over a
hard hollow road, it came in fours with a cadence, a clear yet
hollow sound : it was not long afterwards that I ascertained the
remarkable hammering noise in the air was made by the Jack
Snipe.”
Mr. 8. A. Buturlin, in sending me a specimen of this species,
with characteristic kindness writes the following notes of its habits
as observed on the Kolyma Delta :—“ Its drumming is exceedingly
like the noise of a cantering horse on a hard road, as so well
described by one of the best field-observers—the late John Wolley.
I heard it every day in the summer of 1905, when on the
Kolyma. The bird usually flies so high, that even with the aid
of the midnight sun and good Zeiss binoculars it is often quite
30 MR. P. H. BAHR ON THE [Jan. 15,
invisible; nevertheless the sound ‘ top-toppy-top-toppy * is quite
clearly heard.”
I might also mention here that the remarkable sound produced
by the W ood-Sandpiper (7'otanws glareola) was found by Mr. Buturlin
to be vocal. He shot a score of them ‘“ drumming ” while sitting
on a branch of Ulmus incana, or some local Salix, on the Kolyma,
and has observed them for a long time at the distance of a
few yards. The male flies about in wide circles, beating his wings,
now floating on outstretched wings uttering as loud but not such
hollow notes as the Jack Snipe.
I can only say at present that, in view of the failure to produce
the drumming of the Jack Snipe artificially, I suspect there must
be some other mechanism by which the sound is produced.
Gallinago major: “'The Great Snipe” (text-fig. 8, A).—This
species has 16 tail-feathers, of which the outer four are white. They
are somewhat shorter than the feathers from the centre of the tail,
which are similar in all species of Gallinago. The feathers produce
no sound on experiment. The rami are soft, like those of G. gal-
linula, and can easily be separated. The outer web is composed of
rami provided, as in the case of G@. gallinula, with rows of distal
and proximal radii, of which many are well developed. In the
inner web the distal row is one-third longer than the proximal,
and is provided with 4 feeble hamuli. The rami are inserted into
the shaft at an obtuse angle.
Its breeding-habits have been described by Prof. Collett, of
Christiania, in Dresser’s ‘Birds of Europe,’ vol. vil. p. 635 :—
“The Double Snipe is chiefly a nocturnal bird. Not only does it
migrate at night, but it is in motion almost solely after twilight,
when its peculiar ‘spil* or drumming takes place; and it also
searches after food chiefly during this time of the evening. ...
Tt has a so-called ‘ Leg’ or ‘ Spil,’ like some of the Grouse tribe, a
sort of meeting-place, where they collect to ‘drum’ and often to
engage in combat for the possession of the females. . . . It does not
indulge in aerial evolutions, but remains on the ground. ... The
male bird utters a soft, almost warbling note, which is accompanied
by a peculiar snapping sound caused by striking the mandibles
together several times in quick succession. If a person approaches
one of these drumming-places he can hear at some distance the
low note: ‘bip bip, bipbip, bipbiperere, biperere’; and when
within 100 paces, if the night is still, he begins to hear other
peculiar sounds. ... Whilst producing these notes the bird is in
ecstasy and raises and spreads its tail like a fan, the outer tail-
feathers showing in the halt-darkness like two white patches.”
_ Here, again, having no experience of the aforementioned habits
myself, I can only conjecture that the sound is vocal.
Now we come toa group of Asiatic species, which bear much
resemblance to each other in the structure of their tail-feathers :—
Gallinago solitaria (text-fig. 9, «).—According to the formula
for this species given in Seebohm’s ‘ Geographical Distribution of
1907. | ‘““BLEATING” OF THE SNIPE. 31
the Charadriide,’ this species ought to have 18 tail-feathers ;
but a specimen sent me by Mr. Buturlin from the Western Tian
Shan Mountains has 20 tail-feathers. The outer six are attenuated,
a. Tail of Gallinago solitaria. | b. Tail of G. megala.
the first five markedly so. In this case both the outer and inner
webs are very much narrower. Unlike any of the preceding
species, the rami of the outer web are provided with fully developed
32 MR. P. H. BAHR ON THE [Jan. 15,
vows of distal and proximal radii, the former being provided with
4 hamuli in the same manner as the inner web. Herein this
group differs notably from any of the preceding; the rami are
short and stout, proximal and distal rows of radii the same size.
The bleat produced by these feathers is very loud, and consists
of a number of notes of different pitch intermingled, caused
apparently by the difference in breadth of the different musical
feathers in the tail.
T venture to think that in this group the bleat is produced by
the vibration of the feather as a whole, not by the inner web
alone as in G. calestis.
An account of its breeding-habits is to be found in Hume and
Marshall's ‘Game Birds of India’ :—‘“ They are to be heard and
seen in the higher portion of the hills, soaring to a considerable
height, repeatedly uttering a loud sharp, jerky call, and then
descending rapidly with quivering wings and outspread tail, pro-
ducing a hard buzzing sound, something like, but shriller and
louder than, that produced by G. celestis, though they do not
descend as rapidly as the latter.”
Gallinago megala.—This species inhabits 8.E. Siberia from
Lake Baikal to the North Island of Japan. It possesses 20 tail-
feathers, of which the outermost are 3 inch in diameter. The
attenuated feathers are shorter than those from the middle of
the tail (text-fig. 9, ). Microscopically they resemble those of
G. solitaria in every way. The bleat they produce also resembles
that of the foregoing species, but is far higher in tone. An
account of its breeding-habits is to be found in Taczanowski’s
‘Fauna ornithologique de la Sibérie orientale,’ p. 958, a refer-
ence kindly given me by Prof. Newton. I have translated the
passage from the French: quoting from Prjevalski he says :—
‘That it retires to breed in the deepest marshes, covered with
black scrub, and performs aerial evolutions as follows. The male
soars up in the same way as our Snipe does, and after having
described large circles in its flight above the place in which the
female is nesting, it darts downwards in an oblique direction,
making (probably with the rectrices, as does our Snipe) a loud
sound, like the noise produced by a racquet when the handle has
been broken. ‘This noise gains more and more in intensity as it
approaches the ground, and ceases about 100 paces from it, and
then the bird continues its flight, repeating a note, which one can
express by tic-tic-tic.”
T have received a tail of this species from Central Siberia from
Mr. Buturlin, who sends me this account from his first part of
the ‘ Limicole of Russia’ (1902, pp. 77-78), an account contributed
by the late M. Schwedow :—‘ After 6 o’clock p.m. (in May, near
Irkutsk) one can see in the woods or on forest-swamps the ‘forest
Snipes’ wheeling round and round in the air, and nearly always
repeating in rapid succession notes like ‘chwi, ehwi, chwi,’ or
‘ zswee, zwee, zwee.’ From time to time one or other of the birds
stops beating its wings, somewhat partially closes them, and
swoops obliquely down, while vibrating notes are heard in the
1907. | ‘“‘BLEATING” OF THE SNIPE. 33
air, somewhat like water pouring in the distance, gradually
becoming higher and higher, and more sharply falling on the ear.
The bird falls like an arrow, but when some fathoms from the
tops of the trees it suddenly stops in the air and at the same
moment uttering ‘chic-ka-chee,’ flies on again. After making
some rounds in the air the bird silently or with the same sound
‘zswi, zwsi, goes higher and higher in the air and recommences
the same performance.”
Gallinago stenura (text-fig. 8, C).—The Pin-tailed Snipe of
Tndia breeds in Hast Siberia, and possesses the greatest number of
tail-feathers of the genus—26-28, of which the outer eight or nine
are attenuated. The outer ones are so much attenuated that they
actually do resemble pins. The outer feather measures 545 inch
in diameter, the eighth ;, inch.
The outer web resembles that in G. megala in possessing fully
developed radii; the number of the hamuli in the first pair is 5, in
the eighth pair 4. The ramiare short and thick. These feathers
produce no sound on experiment. The passage from Swinhoe on
Formosan Ornithology, ‘ Ibis, 1863, p. 415, is the quotation from
Taczanowski which I have just given and refers to the preceding
species.
Mr. Buturlin writes of this species on the Kolyma :—‘ It
was only one day, 25th June, 1905, that I could observe its
breeding habits.... I watched it during two or three hours
with strong binoculars at a distance sometimes of not more than
200 yards.... It flew about uttering a high, loud, somewhat
harsh note, not clear enough to be styled a whistle, like ‘ psait,
psait, psait,’ and seemingly produced not by some mechanical
means, but by its voice. From time to time (but not so often as
our Common Snipe) the bird makes head foremost a dive in the
air, just as our Common Snipe, but the descent is in time and
distance quite twice as long as in the common species. When
falling through the air the bird repeats its note more and more
swiftly. ... At the lowest point of its descent, the bird holds the
wings high over its back, just like a swiftly descending pigeon or
duck, and then ascends several feet without evident motion of its
wings. I could not see any opening or spreading out of its tail,
when swooping downwards.”
John Hancock, writing in his ‘ Birds of Northumberland and
Durham,’ found great difficulty in accepting the tail theory,
because of the diversity in structure of the tail-feathers to be met
with in this genus, especially the feathers of G. stenwra, which, he
says, are considered to be musical instruments. Thus he raises
a very reasonable objection, which I shall here quote :—‘ Other
species have the same almost webless feathers at the sides of the
tail, varying only in number. Here then we see a species in
which the so-called sonorous or ‘ musical’ feathers do not possess
the structure, firmness of web, and length of rays, which appear
to be mainly relied on as the sound-producers; though the
rigidity and form of the shaft are in some way or other apparently
Proc. Zoot. Soc.—1907, No. III. 3
34 ON THE ‘‘ BLEATING” OF THE SNIPE. [Jan. 15,
thought to have some influence in the production of the sound,
independently of the rays or web. Were these feathers sonorous
instruments, we should expect to find a greater uniformity in
their structure. But, in fact, the tail-feathers of the true Snipes
are remarkable for their diversity, so much so that the birds have
been divided into four groups, and this mainly on account of a
difference in the number and form of these feathers.”
T have tried to show that the mechanism differs considerably in
different species, just as the sound varies.
For reasons stated before, I believe that the bleat of G. ccelestis
is produced by: the vibration of the inner web as a whole; in
the case of G. frenata, nobilis, and australis by vibrations of the
individual rami; while G. megala and solitaria produce sounds of an
entirely different character by vibration of the feather as a whole.
Finally, I will but briefly mention two species belonging to
closely allied genera :-—
Scolopax rusticola.—The Wopdcock is known to perform certain
evolutions during the breeding-season, producing at the same
time a curious sound, which is acknowledged to be vocal.
Certainly there is no evidence from examining the twelve feathers
of the tail that any of them are specialised structures. The outer
ones do not differ materially either in size or structure from any
of the others. Microscopically the outer web is composed of plain
rami provided with but rudimentary radii. The hamuli are four
in number and their terminal portion is badly hooked.
Philohela minor (Gmel.)—The American Woodcock has 14
tail-feathers, of which the outer ones are decidedly shorter than
the others; they produce no sound in experiment and are in
macroscopical and microscopical structure similar to the last-named
species. A good account of the habits of this bird in the breeding-
season is to be found ih Chapman’s ‘ Birds of Eastern North
America, p. 153 :—-“‘ He begins on the ground with a formal,
periodic, peent peent, an incongruous preparation for the wild
rush that follows. It is repeated several times before he springs
from the ground, and on whistling wings sweeps out the first loop
of a spiral, which may take him 300 feet from the ground. Faster
and faster he goes, louder and shriller sounds his wing song; then,
after a moment’s pause, with darting headlong flight, he pitches
in zigzags to the earth, uttering as he falls a clear, twittering
whistle. He generally returns to near the place from which he
arose, and the peent is at once resumed as a preliminary to another
round in the sky.”
Certain of the primaries of the wing of this species are
characteristically attenuated, for what purpose I am unable to
discover, as they certainly do not produce any sound by any
means I have employed.
T do trust that in the enquiries I have made I may be followed by
1907.] ON THE ANATOMY OF CERTAIN SPECIES Of SQUAMATA. 35
others. I cannot attempt to explain many of the facts I have set
forth in this memoir; and yet an explanation ought to be forth-
coming, and particularly in reference to the microscopical part of
the subject on which I have mainly been able to dwell. The exact
relations of the number and size of the barbules and hamuli to
the sounds they produce is worth investigating, and still more
is the cause of the breaks in the continuity of the sounds which
you have heard—or, rather, not heard. This last would need the
application of one who is intimately acquainted with the science
of acoustics, which I make no pretence to be, and therefore I
cannot offer any suggestion which will account for the non-
continuity of the “ bleating” or ‘ humming”—its sudden stops
and its sudden recurrence. There is much more to be learnt in
this matter, and I would pray those who may be unconvinced by
my experiments, at least to try to account for those marvellous
sounds in some manner more satisfactory, and I assure them that
there is no one who would be better pleased than myself to find
that they can be so accounted for.
In conclusion, I should lke to tender my sincere thanks to
Prof. Newton, of Cambridge, without whose assistance the above
facts would never have been recorded.
Since reading this paper I have received a skin of a female
specimen of G. equatorialis. The sound produced is disappoint-
ing in volume; in tone it bears a resemblance to that of the bleat
of G. calestis.
3. Contributions to the Knowledge of the Systematic
Arrangement and Anatomy of certain Genera and
Species of Squamata. By Frank HE. Bepparp, M.A.,
F.R.S., Prosector to the Society.
[Received December 7, 1906. |
(Text-figures 10-19.)
; CoNnTENTS.
(1) On some Specific Characters of Chameleons shown in the Internal Organs,
p. 3d.
(2) Some Notes upon Chameleolis, p. 45.
(3) The Position of the Umbilicus in certain Vipers, p. 50.
(4) Some Notes upon the Anatomy of Zonwrus, with Special Reference to the
Hyoid, p. 52.
(5) Some new Facts bearing upon the Affinities of Gerrhonotus, p. 56.
(6) On a Point of Structural Resemblance between Heloderma and Varanus,
and on some Specific Characters of Varanus, p. 59.
(1) On some Specific Characters of Chameleons shown in the
Internal Organs.
The external differences among Chameleons are plainly set
forth in vol. iii. of Boulenger’s ‘Catalogue of Lizards in the
a
34
36 MR. F, E, BEDDARD ON THE ANATOMY [Jan. 15,
British Museum.’ The differences between the viscera of different
species are by no means so well known. Indeed the only recent
memoir known to me dealing with the visceral anatomy of
Chameleons, which also refers to specific differences within the
genus, is that by Dr. Wiedersheim*, chiefly dealing with the
respirator y system in Oh. vulgaris and Ch. monachus. 1 have
dissected, with reference to more than one point in the visceral
anatomy, the following species, viz.:—Ch. vulgaris, Ch. calcarifer,
Ch. dilepis, Ch. pumilus, Ch. parvilobus, Ch. teniobronchus, Ch.
basiliscus, and Ch. verrucosus. Iam in consequence able to offer
some additional anatomical facts concerning the genus, which are
also of classificatory importance.
T may first of all call attention to an external character of the
little-known Chameleon calcarifer. One of the external characters
which distinguishes Ch. calearifer from Ch. vulgaris is the presence
in the former of a more distinct ventral crest composed of a
line of conical and, at times, overlapping “enlarged granules.”
This line is traceable, but is by no means so well marked, in
Ch. vulgaris. The division of these ventral scutes which marks
the position of the umbilicus in the foetus is therefore exceedingly
obvious in Ch. calcarifer and less easily to be mapped out in
Ch. vulgaris. It lies behind the middle line of the body. It is
represented by a long space contained in the middle of the ventral
crest, which bifurcates to embrace it, and in this region therefore
1s double. The number of scales on the two sides is uneven ;
T counted 13 on the left and 11 on the right side. The size of the
region of the integument which appears to mark the umbilicus was
rather greater in “Ch. vulgaris, but, as already said, the indistinct-
ness of the ventral crest renders ‘it difficult to be accurate. In
Oh. dilepis this area was quite as distinct as in Ch. calcarifer and
occupied the same position; there were, however, only 10 pairs of
scales, closely apposed.
The lungs show some variation in structure from species to
species. That there is some variation in these organs has already
been pointed out by Wiedersheim, who figures those of Ch. vulgaris
and of Ch. monachus. The differences seem mainly to affect the
number and form of the tags which are appended to the lungs,
those very characteristic anangious outgrowths of the lung.
Wiedersheim distinguishes between the more or less cylindrical
outgrowths and the branches of the lung itself which bear them.
The outgrowths, which exist more anteriorly and always on the
ventral side of the lung, are simply the tubular ceca. Towards
the end of the lung the lung itself is divided into several
processes. In a young example the author quoted found that the
ceca were solid. There is therefore reason for distinguishing the
pulmonary ceca from the lungs, and a careful examination of both
shows a ceasing of the reticular bands which cover even the
anangious part of the lungs. As the number of outgrowths not
* Ber. naturf. Ges. Freiburg-i.-Br., Bd. i. 1886, Heft 3.
1907.] OF CERTAIN SPECIES OF SQUAMATA. 37
only varies in individuals, but also on the two sides of the body
of the same specimen, their mere number and arrangement can
hardly be utilised as distinctive of species without examining a
large series. Yet I am disposed to think, for reasons that will be
discussed later, that Ch. monachus does differ from Ch. vulgaris. |
In Chameleon calcarifer the langs show the same general
structure as do those of Ch. vulgaris. That is to say, the lung
itself is frayed out into processes posteriorly. These again muy
or may not give rise to the tubular cecal outgrowths. The latter
show no network upon their surface, but the direction of the fibres
of which they are partly composed is rather circular. On the
ventral side also, some way in front of the end of the lung, the
lung itself is prolonged into processes. I counted altogether in
one lung examined fifteen tubular cecal outgrowths. But as the
numbers have been stated by Wiedersheim to vary in Ch. vulgaris,
the exact number is probably not a matter of importance. They
were, however, certainly more numerous than in an example of
Oh. vulgaris which I have myself studied. What appears to be
of importance is to note that the lung itself is divided and that
the tubular outgrowths do not arise from a ling with an entire
margin. The subdivisions of the cavity of the lung seem to be
exactly as Wiedersheim has described for Ch. vulgaris. In
Ch. verrucosus the tubular cecal outgrowths are very numerous.
T counted twenty-five or more of them. All the cecal outgrowths
were borne in four tufts, of which that furthest from the bronchus
was the largest, and consisted also of an outgrowth of the lung
itself. The individual tubular ceca were frequently to be seen
arising by the division of a common stem (text-fig. 10, p. 38).
The disposition of the ceca im this species is very different from
that which I have observed in others.
The lungs of Chameleon dilepis appear to differ in certain respects
from those of the species that have been hitherto described. The
obvious difference is the tubular character of the cecal outgrowths,
which have hardly any dilated termination, shown so plainly in
Ch. parvilobus, for example (text-fig. 12, p. 39). In the second
place, the tubular ceeca are thick-walled and not at all transparent
except in parts, and then not so transparent as in other species.
Furthermore, these processes are distinctly shorter in Ch. dilepis
than they are in Ch. parvilobus, as 1s indicated in the annexed figure
(text-fig. 11, p. 39). The differences above set forth can hardly be
due merely to a different state of contraction, since both specimens
came out of the same bottle of aleohol in which they had been
preserved some time since ; and very well preserved, for there was
no trace of softening or disintegration of the viscera. The ceca
of Oh. dilepis are certainly to some extent contracted, as they can
be pulled out without using undue force. There remains, however,
a condition which differs from the attenuated and thin ceca of
Ch. parvilobus and Ch. verrucosus on the one hand, and from the
prolongation of lung-substance with shorter ceca in Ch. calearifer,
‘on the other. The marked distinctness of the czeca from the lung
38 MR. F. E. BEDDARD ON THE ANATOMY [Jan. 15,
is, in fact, a feature of this species as contrasted with those that
have been mentioned. It will be observed in the figure (text-fig. 11)
that the lung terminates in a cecum which continues in the same
straight line. This seems to be the case also with other species. It
suggests, of course, the distal terminal air-sac (abdominal air-sac)
Text-fig. 10.
Lung of Chameleon verrucosus, entire.
of the bird’s lung. The arrangement of the other ceca is shown
in the figure referred to. They are developed along a considerable
region of the ventral margin of the lung. The larger number of
the ceca are, however, massed at one spot, which is not at the end
of the lung as in Oh. parvilobus, but at about its middle. Another
1907, ] OF CERTAIN SPECIES OF SQUAMATA. 39)
noteworthy difference about these ceca as compared with those
of other species which I have examined, is that there is an anasto-
mosis between the roots emerging separately from the lung.
Finally, the small number of czeca as compared, for example, with
Ch. calearifer is a fact worthy of attention, since it is the beginning
of the immense reduction seen in Ch. basiliseus, which culminates
in the total absence of these ceca in Ch. pumilus.
Text-fig. 11. Text-fig. 12.
Text-fig. 11.—Lung of Chameleon dilepis, entire.
Text-fig. 12.—Lung of Chameleon parvilobus, opened longitudinally.
The above description also applies in generalities to the right
lung of the same individual. That is to say, with regard to the
shortness, tubular character, and fewness of the cecal outgrowths.
40 MR. F. E. BEDDARD ON THE ANATOMY [Jan. 15,
A specimen which I had the opportunity of examining fresh
showed how the lungs may vary in individuals, as was pointed
out by Wieder sheim, This variation consists principally in the
Jarger number of ceca. I should mention, however, first of all
that there is just a shadow of a doubt as to the identity of the
species. In the second smaller specimen the occipital lobes
characteristic of the species were disproportionately smaller than
in the larger individual, whose lungs I have already described.
In the second .place, the larger individual had no trace whatever
of the ‘‘ Hohlenvenenfortsatz” of the right lobe of the liver
accompanying the postcaval vein. In the smaller individual there
was a considerable process of hepatic tissue accompanying the
postcaval vein for some distance.
In the case of both right and left lung, the lung ended, in the
same way as in the example already described, in one bifid caecum,
bifid from the very first. In one lung I counted 14 other cecal
outgrowths, of which five were particularly short. In the other
lung I'found as many as 16 outgrowths, of which only three or
four were short. Two, or even three, sometimes borne upon the
same stem. The ceca are of considerable diameter and clubbed
in form ; they contrast markedly with those of Ch. verrucosus.
I have also selected for figuring the lung of Ch. parvilobus (text-
fig. 12), which is at the very opposite extr emity from the other
species figured in the present communication, viz. Ch. dilepis. The
ceca are numerous and extremely slender and in some cases of great
length. Thus the longest measures 33 mm. as against 32 mm. of
the length of the lung itself, and there are several other czca
nearly or quite as long. The longest of the diverticula are at the
posterior end of the lung. The whole ventral border is also beset
with diverticula, but these are invariably short; all show a marked
dilatation at the free extremity. In contrasting these slender ex-.
tended diverticula with the short thick diverticula of the two species
Ch. dilepis and Ch. basiliscus, to be described immediately, one is
disposed to believe that greater contractility in the case of the
two latter may account for the great difference which they show
from the species here under consideration ; especially since Milani’s
figure * of the lung of Ch. basiliscus indicates long slender diverti-
cula with slightly pronounced dilatations at their extremities.
Chameleon basiliscus has lungs which have been described by
Milani? and which agree most nearly perhaps with those of
Ch. dilepis. The lung “itself is extensive and reaches back nearly
to the kidney. The specimen which I had the opportunity of
examining had been preserved for some time in alcohol. In
neither lung could I find any ceca depending from the ventral
margin of the organ, and in the left lung I did not find more
than a single cecum at the posterior end of the lung, but conspi-
cuous enough when detected by its yellow colour as contrasted
with the colourless and transparent wall of the lung itself. In the
* Zool. Jahrb. (Abth. f. Anat.) vil. p. 577.
+ Loe. cit. p. 576.
1907. | OF CERTAIN SPECIES OF SQUAMATA. 4]
right lung, however, though there was a perfect agreement with
the left lung i in the elhsemee of any ventral czecal outgrowths, such
as occur in other Chameleons, there were three obvious ceca at
posterior end. Milani figures five ceca. These were quite tubular
and not swollen at the free extremity, as, for instance, in Ch.
parvilobus. The walls are thick and they arise from a thick-
walled portion of the lung. These ceca are, in fact, exactly like
those of Ch. dilepis, from which the present species mainly differs
in the extreme fewness of the ceca, as 1s apparent from Milani’s
figure.
Chameleon pumilus has lungs which differ in several important
points from those of the species that have been hitherto considered.
In the first place, there are no signs whatever of any bronchi in
the lungs. When the left lung is opened and the appearances
presented compared with those to be seen in Ch. vulgaris, the
following differences are recognisable. In both the aperture of
communication between the two lungs, which represents, of course,
the distal extremity of the bronchus, permits the interior of the
right lung to be to some extent viewed. In the case of Ch. vul-
garis the cartilaginous rings of the bronchus have to be cut up
in order to display fully the aperture into the right lung through
which are seen the cartilaginous rings of the bronchus of the right
lung. In Ch. pumitlus, when the lateral wall of the left lung is
removed no trace whatever of any bronchus is seen; there is
simply a large circular orifice putting the two lungs into com-
munication, which shows no traces of any bronchial cartilages that
can be detected by the unaided eye. It is not plain whether this
condition is to be regarded as primitive or as evidence of degene-
ration. The lung itself is considerably shorter, relatively as well
as actually, than in the species which has been dealt with in the
preceding lines. It is, furthermore, different from the lungs of
these other species in that the typical lung-structure persists
throughout the whole sac. The alveoli in the lungs of Chame-
leons generally are smaller and deeper proximally and get larger
and shallower posteriorly, ultimately becoming pr: actically invisible.
The hinder region of the lung is anangious. In Chameleon pumilus
the alveoli become rather less marked posteriorly, but they are
much more conspicuously circumscribed up to the very end of the
lung than is the case with any of the larger species which I have
had’ the opportunity of examining. The lung, in fact, is less
metamorphosed into a mere air-sac in the present species than in
any other which I have examined, excepting only Ch. tenio-
bronchus, to which species I shall have to refer again immediately.
In this particular it is plain that the lung of Ch. pumilus is more
typically Lacertilian than that of such a species as Ch. vulgaris or
Ch. calcarifer.
A final peculiarity shown by the lung of this species is very
remarkable. It has been stated in many general works that the
Chameleons as a family are to be characterised by the cecal out-
growths of the lungs, which have been considered in several
42 MR. F, E. BEDDARD ON THE ANATOMY [ Jan. 15,
species in the foregoing pages, and that is certainly the general
impression among zoologists and anatomists. I was greatly sur-
prised therefore to find that the lungs of Ch. pwmilus are quite
unprovided with these otherwise characteristic outgrowths. The
margin of the lungs is entire and slightly sinuous, the convexities
occurring in the sinuous line being perhaps to be looked upon as
rudiments or incipia of the cecal appendages. It will be observed
that the absence of these ceca is associated with a more complete
retention of the typical pulmonary structure of the lung, and
therefore its greater efficiency as a breathing-organ. On the other
hand, it is to be noted that where the cecal tubes exist the lung
itself has lost considerably the alveolate structure and thus pre-
sumably some of its efficiency as a breathing-organ. The Ophidia
particularly show that the lung may be too large for its office as
a respiratory organ, and they, like the Chameleons, are often
lethargic in habit.
The above account of the lungs of Chameleon pumilus is, 1 so
far as the absence of tags is concerned, in harmony with the
description of both Meckel* and Cuvier tf. The latter observes :
“ Le Caméléon nain n’a rien de pareil; ses poumons sont deux
petits sacs simples, ovales, de grandeur égale, comme ceux de la
plupart des Sauriens”; and on another page: “ Ils manquent
appendices.” Milani, however, obviously doubts these state-
ments in writing £ as he does: “‘ Ob bei Chameleon pumilus die
Ziptel wirklich fehlen, oder ob diese Behauptung nicht vielleicht
auf ein mangelhaftes Praparat zuriickzufiihren ist, wage ich hier
nicht zu entscheiden.” It is because of the latter doubt cast upon
the facts that I have entered into the matter at some length, and,
as I hope, settled it.
I have finally to add to the description of the lungs in various
Chameleons that Ch. teniobronchus agrees entirely with Ch.
pumilus in the total absence of diverticula, an agreement which is
very significant in view of other facts.
The pigmentation of the interior of the body varies among the
species of this genus. In all that I have examined the intestinal
tract is a deep black, and there are generally (but not in Ch. ver-
rucosus) patches of pigment upon the stomach not distributed so
universally. The mesenteron is largely pigmented anteriorly
in Ch. verrucosus. There is no variation, however, in the pig-
mentation of the gut. The parietal walls are not so generally
pigmented. It is, indeed, only in Ch. pumilus and in Ch. tenio-
brenchus, among the species which I have examined, that the
whole of the lining peritoneum of the body is of a deep black,
quite as deep as is the gut. This pigmentation also extends to
the mesenteries. In all of the remaining species the pigmentation
of the general body-cavity and the mesenteries is hardly to be
seen and only exists in very slight degree, so as not to affect the
* “ Respirationsystem der Reptilien,’ Deutsch. Arch. f. d. Phys. 1818.
+ Lecons @ Anat. Comp. 2me éd. par Duvernoy, t. vil. (Paris, 1840).
ft Zool. Jahrb. (Abth. f. Anat.) vii. p. 573, footnote.
1907.] OF CERTAIN SPECIES OF SQUAMATA. 43
general appearance. This peculiarity at once divides the two
species mentioned from the rest, and other anatomical peculiarities
described in the present communication tend to show the sepa-
rateness of these two Chameleons from others.
It may be remarked that the table of external characters used
by Boulenger in the discrimination of the species of the genus
brings together Ch. pumilus and Ch. teniobronchus*.
Pancreas.—The shape of this organ shows differences in the
species of Chameleon which I have examined. In all it lies
partly between the stomach and the recurrent loop of the duo-
denum, and partly dorsal of the stomach and to the posterior side
of that organ. ‘That is to say, when the reptile is dissected and
viewed in the ordinary position lying on the right side part of the
pancreas, that lying between the stomach and the duodenum is
visible and the rest is seen when the stomach is raised. The main
differences in form are the relative thickness of the gland and the
relations of the splenic lobe, which here, as in other Lizards, is to
be distinguished at least to some extent from the rest of the gland.
The distinction between the two lobes of the pancreas is most
plainly to be observed in Ch. dilepis, where the splenic lobe is
quite at right angles with the rest of the gland, and the duodenal
part is continued on for a very short distance before it gives off
the splenic lobe. In all the remaining species there is no such
marked distinction, the two lobes forming one curved elongated
mass. ‘This is particularly plain in Ch. teniobronchus, where the
coils of the intestine lie entirely behind the pylorus, and the
pancreas is therefore exposed for its whole length and not partially
hidden by the stomach. I shall recur later to the coiling of the
intestine in this and other species of Chameleon.
The bulk of the gland differs greatly in the several species.
In some it is much thinner than in others, and therefore, as the
length is not far from being the same, relatively as to the size of
the species the actual bulk fluctuates. Two extremes are well seen
in the two species Ch, dilepis and Ch. calearifer, which, on account
of their practically identical size, show the facts very plainly. In
Ch. dilepis the gland is very thick, quite as thick as the diameter of
the adjacent pyloric region of the stomach; its greatest diameter
is about 6 mm. On the other hand, in Ch. calcarifer the pancreas
is comparatively quite excessively slender, and only measures
3 mm. or so in transverse diameter in the region which lies
ventrally and in front of the stomach. There are similar differences
between other species; but I do not give details, as the indi-
vidual species vary so much in size that a comparison of the glands
would involve rather complex measurements ; these would be of
more value if the number of individuals examined were large.
The prominent and easily recognisable differences between the
two species selected will serve as an example of what also occurs
elsewhere in the genus. There are, however, too great a series of
* Cat. Lizards Brit. Mus. vol. 11. 1887, p. 440.
A4. MR. F. E, BEDDARD ON THE ANATOMY [Jan. 15,
gradations between the extremes to permit of the use in classi-
fication of the dimensions of this gland.
Liver-lobes—The proportions of the two lobes of the liver *
differ markedly in several species of the Chameleons reported
upon in the present communication. Thus in Ch. pumilus the
right and left lobes are so nearly equal that only one can be seen
with just traces of the other when the viscera are viewed from the
left side. The gall-bladder is partly covered by the extensive left
lobe, which, moreover, comes into contact with the stomach.
The most extremely opposite conditions to these are shown (so
far as the material in my hands enables me to say) in Oh. calearifer.
In that Chameleon the left liver-lobe is very much shorter than
the right. When the animal is viewed in the same posture as the
last, the left lobe leaves exposed a section of the right lobe as long
as itself, and does not even reach the gall-bladder, which lies on
the right lobe not very far from its tip. There is, of course, no
contact between the left lobe of the liver and the stomach, the
ventrally flexed region of which lies considerably behind the end
of even the right liver-lobe.
In Ch, basiliscus the viscera in question are arranged and have
very much the same proportions as in Ch. calearifer. The same
may also be said of the Common Chameleon (Ch. vulgaris) and of
Ch. verrucosus*. In Ch. parvilobus the two lobes of the liver are
approximately equal, and the left lobe completely conceals the
gall-bladder when that viscus is viewed from the left side. On
the other hand, it, the left lobe, does not come so near to the
stomach as in Ch. pumilus. Very much the same description will
serve for Ch. dilepis, save for the fact that in this Chameleon the
gall-bladder is not so completely hidden by the left lobe as in
Ch. parvilobus and Ch. pumilus. The characters of the liver,
therefore, hardly allow of any grouping of the species; for there
are gradations. The alimentary canal does, however, show certain
differences which permit of a grouping such as has been already
suggested.
In Ch. calcarifer and the other larger species the intestine is as
well coiled as in other Lacertilia, and when the animal is opened
from the side a good deal of the small intestine is seen to lie in a
coil with secondary convolutions in front of, 7. e. headward of, the
pyloric end of the stomach. The stomach, in fact, partly covers
a section of the small intestine when the viscera are viewed from
the left side. In Ch. pwmilus there is only one short bend of
duodenum, which lies in front of the stomach. But the opposite
extreme to Ch. calcarifer is to be seen in Ch. teniobronchus. In
this small Chameleon the end of the stomach is not bent upon
itself at all, but is continued back in a straight line to join the
intestine, which is but little coiled upon itself. Moreover, the
whole length of the intestine lies completely behind the stomach.
* The liver itself is very compact in these reptiles, unlobulated, and with very
firm outlines.
+ I could see no gall-bladder in this species.
1907. | OF CERTAIN SPECIES OF SQUAMATA, 45
There is thus a simplification in the coiling of the gut in this very
small species which is not so strongly marked in the rather larger
but still small Ch. pumilus. It is not without interest to note
this apparent relation between smallness of size and simplification
of structure shown also in the lungs of these species, as has been
already commented upon *.
(2) Some Notes upon Chameleolis.
This Lizard is placed among the Iguanide in spite of its super-
ficial likeness to a Chameleon. Indeed this superficial likeness is
not after all very striking, and depends mainly upon the fact that
the head is prolonged behind and above into a parietal crest.
Nevertheless it 1s of advantage to be able to record a few facts in
the visceral anatomy which distinctly confirm the placing of this
genus in the immediate neighbourhood of Jguana. It has, more-
over, its own peculiarities as compared with that genus; and
therefore as a contribution to the visceral anatomy of the Lacer-
tilia I am laying before the Society such facts as I have gathered
from a dissection of a female individual. The existing knowledge
of the anatomy of the Lacertilia shows that there are four
marked structural features in which all the Iguanide that have
been examined agree with each other, and the combination of
which allows them to be defined. I shall, therefore, first of all
deal with these four points, which together prove that Chame-
leolis has been rightly placed among those Lizards.
In the first place, the umbilical hgament which divides the two
liver-lobes and is attached to the ventral median line of the body
is a single ligament which runs continuously from end to end of
the liver, without any trace of a posterior division upon the liver,
as I have lately figured in Jguanat. The gall-bladder is left to
the right of this hgament. In these particulars the umbilical
ligament of Chameleolis is precisely like that of Jgwana, and there
is no need to illustrate the relations of the ligament by a figure.
I may mention that it is deeply pigmented.
A second feature, which, though a small character, appears to be
a constant one, is the position of the intercostal arteries in relation
to the vertebre. In Chameleolis, as in some other Iguanoids,
these arteries plunge into the thickness of the dorsal parietes
towards the posterior end of each vertebra; in some other Lizards
they disappear from view at about the middle of each vertebra.
Of course, Chamcleolis has the same regular arrangement of pairs
of these arteries as in other Lizards, a feature, indeed, which seems
to differentiate the Lacertilia from the Snakes, at least broadly
considered.
Thirdly, Milani £, whose account of the Lacertilian lungs is the
most recent and comprehensive known to me, has found that in
the Iguanide the lung is totally divided into two chambers, of
* Supra, pp. 39 & 41.
+ P.Z.S. 1905, vol. i. p. 12, fig. 7.
{ Zool. Jahrb. (Abth. f. Anat.) vii. p. 545.
46 MR. F. E, BEDDARD ON THE ANATOMY [Jan. 15,
which the more dorsal extends headwards of the orifice of the
bronchus. This statement at any rate holds good for the majority
of the Iguanide that have been examined. Chameleolis does not
agree with these types, for the lung is not divided by an obliquely
placed septum into two approximately parallel chambers. It is,
nevertheless, not to be removed from the Iguanide on this
account, since it appears to present points of likeness to the
undoubtedly Squamoid Phrynosoma, and one point at least of
resemblance to the Iguanoid genus Polychrus.
Text-fig. 13.
Lung of Chameleolis, opened longitudinally.
The bronchus enters for a short distance into the lung as a
completely circular tube; there is no snake-like series of flattened
semirings such as is to be found in Jguana*. The projecting
bronchus is, as in Phrynosoma, moored to the walls of the lung by
septa. The cavity of the lung, therefore, extends headwards of
the opening of the bronchus and all round it. There is no septum
in either lung which separates off the dorsally placed caecum of the
lung as a distinct cavity from the rest of the cavity of the organ.
In the left lung the structure happened to be more favourable for
observation than the right lung, and I have accordingly had a
drawing prepared (text-fig. 13) of the interior of this lung. It
* Milani, Joc. cit. pl. xxxi. fig. 13.
1907. | OF CERTAIN SPECIES OF SQUAMATA. 47
will be there seen that the strong septa which produce a pouching
of the dorsal region of the lung in other Iguanids and Agamids
are also to be seen in Chameleolis. Lobserved six of the chambers
altogether, of which three would appear to belong to the anterior
part of the lung, 7. e., that region which is in other Iguanids
divided off by a septum from the posterior region, and three larger
pouches belonging to the posterior region of the lung. Finally,
the end of the lung abruptly narrows and forms a finger-shaped
region with a but slightly marked network. It seems to me to be
possible to compare this with the Chameleon-like outgrowths of
the lung in Polychrus marmoratus *.
In the fourth place, the right extremity of the liver is attached
by a fold of membrane which separates the lung from the post-
hepatic region of the body-cavity and is continuous with the
oviducal membrane.
Besides these points, which, together with various external and
osteological characters used by others, fix the systematic position
of Chameleolis, there are other features in its anatomy which I
have ascertained and which are worth noting as a contribution to
Lacertilian structure.
The pigmentation of the body-cavity is in some ways remark-
able. The umbilical ligament, not only the region which is
attached to the liver, but that which is attached to the stomach,
is deep black, and in the latter region contrasts with the yellowish
gut. The gut itself is, however, pigmented in the case of the
large intestine. This pigmentation is limited to the dorsal side
of the gut and involves the whole of the cecum, The appearance
presented is, indeed, of two tubes closely applied, of which one is
the small intestine and the other ends at the blind extremity of
the cecum. _
As in many Lacertilia, the peritoneum generally is deeply pig-
mented, and a distinction is to be drawn between the posterior
pigmented region and the anterior region of the body-cavity,
where its walls are not pigmented at all, so far as naked-eye
appearances go.
While, however, in most Lacertilia this line of demarcation is
quite oblique, bending ventrally in a continuous curve, it is in
Chameleolis quite transverse (to the longitudinal axis of the body)
in direction, but with a curved outline, now convex, now concave.
The existence of bundles of plein muscular fibres in the mes-
enteries reaches a very great degree of development in many
Lacertilia. In the Lizard which forms the subject of the present
communication there was no development of such fibres that could
be seen with the unaided eye. The ovaries contained no mature
ova. There was a fully formed egg in each oviduct, with a dirty
white shell of leathery consistency. There was no trace of an
embryo in the egg.
The apex of the heart is fixed to the pericardium by a very
* Milani, loc. cit. pl. xxxi. fig. 15,
48 MR. F, E. BEDDARD ON THE ANATOMY [Jan. 15,
slender gubernaculum cordis, a structure which is rarely absent
from the heart of the Lacertilia *.
The arterial system presents certain peculiarities as compared
with that of other Lacertilia. The disposition of the intercostal
Text-fig. 14.
B ~~ -Coel.
A. Stomach and pancreas of Chameleolis.
P. Pancreas; p.v. Portal vein; p.v.g. Gastric portals; Sp. Spleen.
B. Dorsal aorta and branches of the same.
Cel. Coeliac artery ; i. Intestinal arteries; ws. @sophageal arteries ; Sp. Spleen.
arteries has alveady been mentioned as a point of affinity with the
Iguanide. The visceral arteries (text-fig. 14) which supply the
* But is absent m Varanus occasionally (see Beddard, P. Z.S. 1906, vol. ii. p. 617
footnote) and Zonurus giganteus (infra, p. 55).
1907. | OF CERTAIN SPECIES OF SQUAMATA, 49
alimentary canal are collected into two groups, leaving a tract of
considerable length from which no arteries to the gut arise. The
anterior group is situated just behind the union of the two aortic
arches, and consists of no less than seven small arteries supplying
the cesophagus and stomach, ‘These arise from both sides of the
aorta and are partly arranged in pairs; they run to both sides of
the stomach, There is then a long gap until the origin of the
intestinal arteries. The general ‘plan of these is like that in
most Lacertilia ; but there are differences in detail from those of
many genera, When the mesentery is turned over to the right
the cecal arte xy which arises most anteriorly is seen to run over
the following duodenal artery, but under the third artery, that
which : supplies the spleen, &e,
With regard to the venous system, the only notes that I have
made refer to the hepatic por tal system. The junction of the
anterior abdominal and the main portal trunk is very near to the
conjoint entrance of both into the liver—much nearer than is the
case with many Lizards. In addition to this, the chief portal
trunk, there are two vessels which pour blood direct from the
stomach into the liver (text-fig. 14): one of these, the more
posterior in position and the larger, is associated with the left
lobe; the other, a slender twig, enters the liver to the right of the
last described. The ventral parietal hepatics are also two, of which
one is a little to the right of the other in its point of entrance to
the liver. Both are rather far forward on the liver. There is
but one dorsal parieto-hepatic. This, as is the case with other
Lizards*, is associated with the “ Hohlvenenfortsatz” of the liver,
and runs in the mesentery, binding that lobe of the liver to
the right parietes. It runs a conside ‘able way forwards along
the ver rtebral column before becoming lost in the thickness of the
parietes.
The pancreas of Chameleolis (text-fig. 14) i is constructed upon
the usual Lacertilian plan, but differs in various details from that
of other Lizards. It isa Y-shapel gland and completely solid
throughout. There are no thin diffuse branches spread through
the mesentery such as are to be found in the case of the pancreas
of Zonurus giganteust. One arm of the ¥ ends, after dilating
slightly, in the concavity of the somewhat bean- shaped spleen ; the
other forms a thick mass in contact with the commencement of
the duodenum. The stem of the Y forms a thin rod of pancreatic
tissue, which closely accompanies the portal vein and very nearly
touches the liver. This region of the pancreas seems to me to be
longer than in some other Lacertilia, though in most there is a
process of the pancreas running in the same direction. The
splenic lobe of the pancreas is not extraordinarily thin, as it is in
Tiliqua scincoides ~, but of fairly robust diameter,
* The absence of this vessel is rare, but Hochstetter, whom I have been able to
confirm, has asserted its absence in Chameleon vulgaris. I take this opportunity
of stating that this vein is also absent 1 in Chameleon verrucosus.
+ Vide infra, p. 55. { See Beddard, P. Z. §. 1905, vol. ii. p. 262
Proc. Zoou. Soc.—1907, No. IV. 4
50) MR. F, E, BEDDARD ON THE ANATOMY (Jam. 15,
(3) The Position of the Umbilicus in certain Vipers.
IT am not aware that the point of entrance of the umbilical sac
into the body in Snakes has ever been made use of as a systematic
character. I find, however, from a few observations that I have
been able to make recently, that this anatomical relationship is
apparently of systematic value. Since of one species, viz. Lachesis
lanceolatus, selected for these observations, I have been able to
examine a considerable number of individuals, the variation of
the character from one individual to another became a matter of
additional interest, especially in view of the fact that all the indi-
viduals were of one brood. It appears that in Vipers, as compared
at any rate with the Anaconda*, the umbilicus is much nearer tothe
cloacal aperture. I have examined fourteen individuals of Lachesis
lanceolatus of the same brood and of approximately the same size,
though they died on different dates, from March the 9th to May.
The length varied from 113 to 12 inches exclusive of the short
tail. I do not give measurements in millimetres, since to use such
gives an appearance of rigid accuracy not attainable in a dead
snake capable of artificial extension and shortening. In nearly
all of these fourteen individuals four scales occupied the umbilical
region, each of them being bisected by a groove running longi-
tudinally to the axis of the body. I found, in fact, that there
were four scales thus modified in eleven individuals. In two of
the remaining snakes there were five of these scales in which the
two sides had not joined across the middle ventral line, and in
the fourteenth individual only three scales and a portion of the
fourth ; the number of scales intervening between the last of the
‘‘ umbilical” scales and the anal scale varied a good deal but within
very narrow limits. The actual facts are these: in three speci-
mens 17 scales intervened between the points mentioned ; in one
specimen 18 scales; in five others 19 scales; in three 20 scales;
in one 21 scales; and, finally, in one 22 scales. The average is
thus arithmetically 19, and actually there were more specimens
exhibiting the average than any other number. Having due
regard to the narrow.range of the variation, it seems likely that
the position of the umbilicus in this species of Viper can be
regarded with safety as lying 19 scales in front of the anal scale.
It is important to notice the length of time during which this
feetal character is retained. The last specimens examined by
myself died on May 15th of last year. These and the other
individuals were acquired by the Society on Dec. 12th, 1905.
The last specimens examined by me were therefore more than six
months old. J have some confidence, therefore, in comparing
Lachesis lanceolatus in respect of these characters with other
Vipers of an obviously greater age. I may first of all, however,
refer to newly-born Vipers which I have recently dealt with tf in
* See Beddard, P. Z.S. 1906, vol. i. p. 13.
+ P.Z.S. 1906, vol. i. p. 34.
1907. | OF CERTAIN SPECIES OF SQUAMATA, 51
a paper communicated to this Society. It is impossible to be
certain of the exact position of the actual umbilicus in Lachesis
lanceolatus for the purposes of comparison between these two
types, 7. e., which of the four or five broken scales correspond to
the two scales in Bitis nasicornis which are actually divided by
the foetal blood-vessels. Assuming, however, that they are even
the last two, there still remains a substantial difference in position
between the umbilicus of the twospecies. For in Ditis nasicornis
the actual numbers of scales intervening between the umbilicus
and the anal scale are respectively in the five examples studied 9,
11, 11,12,14. There is thus exactly the same amount of variation
as in Lachesis lanceolatus, but round a different mean.
The position of the umbilicus in Russell’s Viper (Vipera
russellit) is again different and relatively more fixed than in either
of the species hitherto considered. As in Sitis nasicornis, the
actual umbilicus consists of two scales only, which do not meet
ventrally, and between which the plug of tissue bearing the
umbilical vein &e. passes into the interior of the body. The
young Vipers in question were a very few days old, but all
external traces of the yolk-sac had disappeared. In front of
these two scales either two or three scales were divided by a suture
in the middle ventral line, and posteriorly to the two ‘“ umbilical
scales ” either one or two scales were similarly split by a ventral
suture. Between the last of the two completely divided scales
and the anal scale there intervene in the five examples examined
respectively 16, 16, 16, 17, 17 scales. ‘The position of the umbi-
licus is therefore different in this Viper, and its fluctuations of
position are less than in the two species to which I have already
called attention. It is perhaps permissible to call attention to
the fact that Vipera and Lachesis agree with one another more
nearly than either does with Sitis. This is, of course, not in
accord with generally received views upon the classification of
Vipers.
J have examined several Vipers of more mature age, and in
two specimens, at any rate, I find what appear to be obvious traces
of the umbilicus. In a not fully-grown example of itis arietans
measuring 30 inches from the snout to the cloaca, four scales
showed a line of division in the ventral median line. The second
of these had the most strongly marked groove, and_ possibly
therefore represents one of the two scales already described in
the young as immediately surrounding the stalk of the yolk-sac.
Between the last of the grooved scales and the anal scale 9 scales
intervened. ‘The species evidently therefore comes nearest to the
other species of Sitis which has been described above. Inasecond
specimen measuring 32 inches there were 12 scales between the
last of four grooved scales and the anal scale. I have also seen
similar traces in a large adult example of Bitis gabonica. Here
there were also four scales showing traces of the umbilicus; but
instead of being grooved they were merely nicked posteriorly.
Between the last of them and the anal scale 8 scales interyened
4?
52 MR. F. E. BEDDARD ON THE ANATOMY [Jan. 15,
Tt is not safe upon these two last-mentioned examples to attempt
to draw any distinctions between the different species of Bitis.
It seems, however, to be most probable that they do not differ
widely from each other as regards the points under discussion,
whether they will ultimately be found to differ specifically or not.
It is, however, quite plain, in reviewing all the facts brought
forward in the present communication, that the position of the
umbilicus among Vipers is one that does at least characterise
some forms which happen in the instances studied to be generi-
cally separated.
(4) Some Notes upon the Anatomy of Zonurus, with Special
Reference to the Hyoid.
The following notes refer to three specimens of Zonurus
giganteus which I have had the opportunity of dissecting during
the last year or two. The anatomy of this Lizard is already to
some extent known through the work of previous observers.
The lungs have been dealt with by Milani* in his general account
of these organs among the Lacertilia, and the arteries of the
gastric and intestinal regions are described and figured by Hoch-
stetter +. There remain, however, a few points to which it is
worth while calling attention as a further contribution to the
natural history of this Lacertilian.
Of special importance—rather, however, from a general point
of view than as a particular contribution to our knowledge of this
Lizard—is the condition of the elements which together make up
the hyoid complex of bones and cartilages in this Lacertilian.
T am able to add to what I have to say concerning Zonurus a few
notes upon other genera of Lacertilia which I have dissected for
purposes of comparison. I commence with a brief résumé of
some of the facts already known of this part of the skeleton.
The hyoid and branchial arches of Lacertilia have not, as it
appears, been investigated in a very large number of genera.
Several are figured in the volumet of Bronn’s ‘'Thierreich’ dealing
with the Lacertilia, while other genera have been illustrated by
subsequent writers §. Apart from differences in the form of the
individual elements of the hyoid complex there is substantial
agreement, according to these various writers. For contrary to
what is to be found in the Chelonia—where the remains of the
hyoid arch proper is followed by two branchial bars considerably
developed—the Lacertilia are generally believed to be characterised
by the preservation in the adult of only one visceral arch follow-
ing the hyoid arch, which is stated to be the first branchial.
This statement occurs at any rate in such authoritative textbooks
* Zool. Jahrb. (Abth. f. Anat.) vil. p. 545.
+ Morph. Jahrb. xxvi.
+ Reptilia, Bd. vi. Abth. iii. Taf. 72. figs. 2-8, & Taf. 107. figs. 24, 33.
§ E. g., Gecko mauritanicus, Gadow, Phil. Trans. 1888 B, pl. 72. fig. 10; Helo-
derma suspectum, Shufeldt, P.Z.S. 1890, pl. xviii. fig. 6; Chlamydosaurus and
Physignathus, Beddard, P. Z. 8. 1908, vol. 1. p. 20, text-fig. 9,and p. 21, text-fig. 10.
1907. | OF CERTAIN SPECIES OF SQUAMATA. 53
as those of Hatchett Jackson *, Gadow 7, and Sedgwick ¢, which
may be regarded as expressing the current knowledge of the
subject. Nevertheless, the late Prof. W. K. Parker$, in describing
the adult skull of Lacerta agilis, wrote as follows :—‘‘ Another bar,
half as long as the first, and unossified, lies behind the first
branchial above; it is f-shaped, with the top hooked inwards,
like the lower piece; this is the upper (0r.?), or ‘epibranchial’
part ; it has a small snag outside its middle. Besides this, there
is, on each side, a slender, slightly outbent hypo-branchial (/.67r.);
this belongs to the second branchial, and also from its length is
evidently part of the third, neither of which chondrify, above, in
the embryo.” In a footnote is added the remark that “this little
highly-metamorphosed Lizard has scarcely thrown aside the
skeleton of these organs of aquatic respiration.” It is obvious
that Prof. Parker’s account is a little misleading, and this doubt-
less accounts for the fact that the existence of remains of a second
branchial arch in Lacerta has been largely ignored in zoological
literature. What he speaks of as an “ epibranchial,” without
determining to which arch it belongs, but letters “ br.*,” is clearly
from its position a vestige of the second branchial arch, as is plainly
recognised in Prof. T. J. Parker’s ‘Zootomy’ and in his ‘Textbook
of Zoology,’ written in conjunction with Prof. Haswell|. The
exceptional character of the hyoid complex of Lacerta in possess-
ing “the epibranchial of the second branchial arch” is properly
emphasised by Dr. Shufeldt {| in reviewing existing knowledge of
the Lacertilian hyoid bones.
The third postmandibular arch is, however, by no means a
peculiarity restricted to Laverta. It occurs in Zonurus in the
form of a short and slender bar lying behind the well-developed
first branchial bar **. This bar of cartilage does not extend down
to the median copula, and indeed falls a considerable distance
short of it. I have examined three other Lacertilians in which
this same visceral arch is represented and one in which it is not
to be found; but I have not at present attempted an exhaustive
research into the facts of its absence or presence among the
different families of Lacertilia. I could not detect the bar of
cartilage in Chameleolis, whose anatomy has been described above.
Tt is well developed in both 7iliqua and Trachydosaurus. Inthe
former (text-fig. 15, p. 54) it is very conspicuous, and it is nota
little surprising that it has been missed, unless I have unwittingly
overlooked its description somewhere. But if this be the case it is
clear that its existence has escaped the writers of many textbooks.
* Ond ed. of Rolleston’s ‘ Forms of Animal Life.’
+ “ Amphibia and Reptilia,” in ‘Cambridge Natural History.’
t ‘Textbook of Zoology,’ vol. 1.
§ “Development of Skull in Lacertilia,’ Phil. Trans. 1879, p. 616.
|| ‘Textbook of Zoology,’ vol. 11.
q P.Z.S. 1890, p. 225.
*k There is no trace of this shown in a figure of the hyoid of Zonurus cordylus
copied from Henle in Bronn’s ‘ Thierreich,’ Reptilien, vol. vi. Abth. 11. Taf. 107.
fig. 33.
54 MR. F, E. BEDDARD ON THE ANATOMY (Jan. 15,
In Tiliqua this second branchial arch is more extensive, as it
appears, than in Zonurus, and lies obliquely across the first
branchial arch, though beneath it. The latter arch ends in a
curled piece of cartilage which is directed backwards and overlaps
the third post-mandibular arch. But the position varies according
to the degree of distortion of the muscles of the neck of the reptile.
Text-fig. 15.
H Br.
Tiliqua scincoides, head and neck.
Dissected to show three postmandibular visceral arches in sitw.
To the left the isolated extremities of the same arches in another individual.
Br.1. First branchial arch; Br.2. Second branchial arch ; H. Hyoid arch.
Towards the lower end of the bar is a triangular, projecting,
“snag” (not visible in one of two examples dissected) like that which
Prof. W. K. Parker has figured in Lacerta. To this projecting
process is fixed a ligament which is inserted on the free dorsal end
1907. | OF CERTAIN SPECIES OF SQUAMATA. 59
of the hyoid arch. The ligament thus avoids the first branchial
under which it lies. It is not surprising to find that 7rachydo-
saurus, so closely allied to Z'iliqua, also possesses this second
branchial arch. As in the genera mentioned, the arch is only
represented by its upper part, the epibranchial, as Parker termed
the equivalent cartilage in Lacerta. Finally, I have to record
that the bar of cartilage is also found in Gerrhonotus, a genus
of whose anatomy I offer some further notes below. In the
present state of our knowledge it is not possible to state whether
or not this occurrence does or does not bear wpon the affinities of
Gerrhonotus. The cartilage was not so easy to find in this small
Lizard, where it is siender and delicate, but can be detected
by gently moving in various directions the muscles in its vicinity ;
the stiff ends of the cartilage thus become apparent.
A second feature in the anatomy of Zonwrus to which I desire
to draw attention is the total absence of the gubernaculum fixing
the apex of the ventricle to the walls of the pericardium. This
ligamentous band or thread (it varies in importance in different
genera) is so usual among the Lacertilia as to be characteristic of
that order of Reptiles, as it is, indeed, of others. I have already
pointed out that the gubernaculum cordis is not to be found in
the heart of Varanus niloticus and some other species*. It is
interesting to notice that this absence of the gubernaculum which
is universal in the higher Vertebrates (Aves and Mammalia), as
well as generally in the Ophidia *, is sporadically developed among
the Lacertilia. It should also be mentioned that this condition
of the heart was found in two examples of Zonurus giganteus (the
third was not examined ad hoc)t; it is therefore probably charac-
teristic of the species if not of the genus.
The liver of this Lizard is unusual in its form. The right lobe
is prolonged in the usual way over the vena cava. But the left
lobe, instead of being but slightly divided at the entry of the
anterior abdominal vein, is deeply bifid thereat §. The whole
organ is thus markedly trifid posteriorly and is not unsuggestive,
in appearance, of the mammalian liver.
The pancreas displays one noteworthy character. Its general
form is like that of the majority of Lacertilia. The organ embraces
the stomach, being found on both sides of it; the splenic lobe is
fairly stout and reaches the spleen, and there is a process of the
gland extending towards the liver. The peculiarity of the pancreas
of this Lizard is that diffuse thin ramifying tags of pancreatic
tissue lie in the mesentery on either side of the splenic lobe of the
gland with which they are connected. This tendency towards a
diffuse irregularly shaped thin pancreas is obviously to be com-
pared with the conditions obtaining in the Chelonia.
* P, Z.S. 1906, vol. ii. p. 617 footnote.
+ Perhaps universally also. In any case the occasional ligament tying the apex
of the heart to the pericardium is rather different (see P. Z.S. 1904, vol. ii. p. 107).
+ I have since found the same absence of the ligament in another example.
§ I am not quite certain that it is not the right lobe which is thus bifid. It is a
point difficult to settle.
56 MR. F, E. BEDDARD ON THE ANATOMY [Jan. 15,
(5) Some new Facts bearing upon the Affinities of
Gerrhonotus.
This genus of Lacertilia is sometimes placed in a special family
of Lacertilia, the Gerrhonotide. By others (¢. g., Boulenger *,
Gadow *) it is relegated to the Anguide. The general aspect otf
Gerrhonotus ceruleus is, on the other hand, by no means unlike
that of the Scincide ; I am not aware that any notes have ever
been published upon the visceral anatomy of this Lizard. I
venture, therefore, to lay before the Society some notes which a
recent dissection of more than one example of Gerrhonotus ceruleus
enables me to offer as a contribution towards the determination
of the systematic position of this genus or representative of a
family.
T have by no means attempted an exhaustive survey of the
anatomy of this Lizard. But J] am able to note down a few facts,
all of which are of some interest from the point of view of a
comparison with other Lacertilia. The structure of the guadrato-
jugal ligament is one of the characters which I carefully examined
in Gerrhonotus. I find that the arrangement and appearance of
this ligament is precisely as it is in the genus Gerrhosaurus and
in the Skink Humeces~. That is to say, the ligament of this
genus is very distinctly marked off and of equal breadth through-
out, nowhere vaguely shading off into surrounding tissues.
Moreover, it is attached on the one hand, of course, to the quadrate
bone and on the other to the bony scales which cover the face in
this region. It is not inserted on to any bone of the skull. In
the present state of our knowledge it is not possible to comment
upon this likeness to Gerrhosaurus and Hwmeces as an argument
in favour of the Skinkoid affinities of Gerrhonotus, though I have
thought it worth while to record the fact for future comparison.
The second feature in the structure to which I draw attention is the
complete pigmentation of the interior of the body. There is here
no paler area divided by the oviducal mesentery from a more
darkly pigmented posterior portion.
As is now well known §, the umbilical hgament of the Skinks
is frequently a double ligament attached to the ventral surface
of the liver along two parallel lines which become confluent
anteriorly. I observed nothing of the kind in Gerrhonotus, where
the umbilical ligament is, as in most Lizards, a single mesentery.
Tn this anatomical fact there is a likeness to Ophiosaurus as well
as, of course, to Zgwanaand other Lizards. In any case the Lizard
shows no affinities to the Scincide.
The pancreas and the spleen and their relationship to one
another differ greatly among the Lacertilia, and more than one
fact in the structure of the two glands is recorded in the present
* Catalogue of Lizards in the Collection of the British Museum.
+ “Reptilia,” in ‘Cambridge Natural History,’ p. 538.
t P. Z.S. 1905, vol. ii. p. 256.
§ Beddard, P. Z.S. 1888, p. 102.
1907, ] OF CERTAIN SPECIES OF SQUAMATA, 57
communication *, In Gerrhonotus the splenic prolongation of the
pancreas is present, but it does not reach the spleen at all, though
extending a good way in the direction of that organ. Among
the Skinks this pancreatic process towards the spleen is to be
found, as I have already recorded‘, in the genus J%liqua, and can
confirm in all details from a subsequently examined example of
that genus.
There is, however, no particular likeness in the structure of the
pancreas of Gerrhonotus to that of Ophisaurust. In the latter
the pancreas consists only of two closely applied lobes which rest
upon the ventral surface of the pylorus and small intestine, there
is no vestige of a splenic lobe§. The spleen of Gerrhonotus is
rather peculiar in position. Very generally among the Lacertilia
this blood-gland is elongated and somewhat bean-shaped in out-
line, and hes with its long axis parallel with the long axis of the
stomach. In Gerrhonotus the shape is quite normal, but the long
axis 18 perpendicular to the long axis of the stomach.
The hepatic portal system of veins of Gerrhonotus ceruleus
varied but little in the two specimens dissected. The ventral
parieto-hepatic veins running in the umbilical ligament were
three in each Lizard. The first two crossed each in their course
in one specimen, and perhaps in both, though I have no note as
to this in the second example dissected. ‘The crossing is such
that the anterior of the two vessels draws blood from a region of
the ventral body behind that which is supplied by the posterior
of the two veins.
The dorsal parieto-hepatics are either two or three and are
otherwise quite normal in their position. The arrangement of
the gastro-hepatic veins is interesting in relation to the question
of the affinities of the genus Gerrhonotus. There are either four
or five of these vessels of somewhat varying calibre arranged close
together, and thus forming a ladder-like structure lying quite at
the anterior end of the liver and running to this from the adjacent
border of the stomach. There are no gastro-hepatic veins situated
more posteriorly. The interest attaching to this arrangement of
the vessels is that it is completely paralleled in Ophisawrus apus ||,
making allowances for the greater elongation of the liver in the
latter snake-like Lizard.
In Ophisaurus, in fact, there are six of these veins. Now, as a
rule, the Lacertilia have not a great many separate gastro-hepatic
vessels. J have myself examined several species embracing as
many genera and find the following facts, some of which I have
* Cf. pp. 48 & 55. + P.Z.S. 1905, vol. ii. p. 262.
{ In contrast to this difference in form between the pancreas of genera which
appear to be allied is the close resemblance in another case which I take this oppor-
tunity of recording. In both Iguana tuberculata and Liolemus magellanicus (I
owe the specimen to the kindness of my friend Capt. Richard Crawshay), which are
both Iguanide, but not much alike superficially, the long splenic lobe of the pancreas
just touches the posterior end of the spleen.
§ See P. Z.S. 1905, vol. ii. p. 4.75, text-fig. 64. Anguis also lacks the splenic
lobe.
|| See P. Z. S. 1905, vol. ii. p. 475, text-fig. 64.
58 MR. F. E. BEDDARD ON THE ANATOMY [Jan. 15,
already made known in recent communications to the ponely
upon the anatomy of these Reptiles. In Jgwana tuberculata I
found in two examples two gastro-hepatic veins, and precisely the
same arrangement characterised two examples of Amphibolurus
barbatus. Uromastix acanthinurus showed, also in two examples,
a single vein, which, however, was made up of three considerable
affluents from the stomach ; these, it will be understood, entered
the liver as a single vessel. In one of the specimens the third
affluent only joined the common trunk formed by the other two
just before their entrance into the liver. In Gerrhonotus, Tupi-
nambis, Chameleon, Phelsuma, Tarentola, | have recorded, or am
now able to record, the existence of only one gastro-hepatic vein,
which however is, as a rule, made up of two affluents. The
Scincide form an exception to the general arrangement of these
vessels, and at first sight appear therefore to be near akin in this
particular to Gerrhonotus.
Of Tiliqua scincoides | have dissected two examples for the
purposes of the present investigation, and find in both of them
the following arrangement of the gastro-hepatic veins. There
are four of these, which appear at first sight to lie accurately side
by side in the gastro-hepatic mesentery. A more careful exami-
nation, however, shows that the stomach is bound to the liver by
two mesenteries, one above the other, as seen when the animal is
opened along the median ventral line and the viscera examined
in an tndissmbed condition. The lower of these, i.e. that which
lies above in the ordinary position adopted in dissection, is the
gastro-hepatic mesentery found in all Lizards. When this is cut
through a second mesentery comes into view, which is attached to
the right side of the liver and to the more dorsal side of the
stomach. This mesentery exists in other Saurians, but is inserted
on to the mesogastrium and does not touch the stomach at all.
Whether this arrangement of the right dorsal suspensory hgament
of the liver has anything to do with the double umbilical hgament
of the same family of Lizards is not certain; but it is found in
most but not in all Skinks. I find it in the genus which we are
now considering, in Seps (Chalcides), Scineus, Hwmeces, and Macro-
scincus. It is not to be found in Trachydosaurus rugosus. To
revert to the gastro-hepatic veims in Z'%liqua scincoides, the most
posterior of the four veins runs along the right or lower (as seen
on dissection) gastro-hepatic mesentery; in front of it are two
of the veins which run in the upper or left gastro-hepatic mesen-
tery (the mesentery present in all Lizards). Finally, there is a
single vein which is inserted just at the junction anteriorly of the
lines of attachment of the two mesenter ies, to the lower (dorsal)
surface of the liver. In Trach ydasourus, which, although a
member of the family Scincide, agrees with other Lizards in the
presence of only one gastro-hepatic mesentery, I find in an
example dissected only » one gastro-hepatic vein, which, as is so
usual, is formed by two equally sized affluents, I have some
notes, however, of an example, dissected a good many years ago,
e
1907. | OF CERTAIN SPECIES OF SQUAMATA. 59
in which in addition to this there was another vein further
forward at the junction of the two dorsal suspensory mesenteries
of the liver. dMJacroscincus is normal—for a Skink—ain the presence
of two gastro-hepatic mesenteries ; and yet it has only one gastro-
hepatic vein. This is formed of two equisized affluents, and runs
in the right-hand mesentery, the other being quite anangious.
Eumeces algeriensis shows the same double series of gastro-
hepatic veins that are to be found in Z%liqua scincoides. ‘There
is one vein only in each of the two gastro-hepatic hgaments and
a third vein implanted at the junction of these anteriorly. As
in Tiliqua, the medianly situate vein of the three belongs to the
left-hand ligament. Of an example of Chalcides ocellatus, dissected
by me a good many years ago, I have sketches ae descriptions
showing that this species is more like Macroscincus* than Humeces
or Piligua. For the gastro-hepatic veins are limited to the right-
hand one of the two gastro-hepatic ligaments, with the usual
vein which enters the liver at the junction of the two veins.
There were in this individual four of these veins in the right
ligament. It seems, therefore, that the numerous gastvo-hepatic
veins of the genus Gerrhonotus may be regarded as evidence of
affinity with the Anguide, since these veins are, as a rule, not
numerous among the Lacertilia other than the Anguide, Amphis-
beenidee, and arent “ia, except among the cemenees where the
existence of two castro- hepatic ligaments accounts for the greater
number of these veins than occurs in the majority of those genera
with only one gastro-hepatic ligament. And, in coming to this
conclusion, it must be further borne in mind that the number of
gastro-hepatic veins would appear to be fairly constant, so far as
the somewhat meagre facts already known allow us to judge.
There is but little alse in the anatomy of this Lizard, so far as I
have been able to record the facts, which bears very distinctly
upon its affinities, a conclusion for which the, at present, very
small knowledge of the Lacertilia is doubtless largely responsible.
It seems, at any rate, to be the fact that Gerrhonotus exhibits no
marked features in its organisation which poimt to an affinity
with the Scincide, except perhaps the condition of the quadrato-
jugal ligament, which is undoubtedly like that of Huwmeces.
(6) On a Point of Structural Resemblance between Heloderma
and Varanus, and on some Specific Characters of Varvanus.
Although it is not the prevalent opinion that these two genera
of Saurians are nearly allied, there are not wanting anatomical
resemblances between them; and, indeed, some of the most recent
writers 7 on the anatomy of Heloderma have brought together a
* It is perhaps not without interest to notice that in these two genera (Chalcides
and Macroscineus) the double character of the umbilical ligament is not so marked
as it is in Humeces and Tiliqua. They have, therefore, at any rate, two anatomical
features in common.
+ Boulenger, P.Z.S. 1891, p. 109; Beddard, ibid. 1906, vol. ii. p. 601.
a
60 MR. F. E, BEDDARD ON THE ANATOMY [Jan. 15,
good many facts in favour of such an alliance, On the other
hand, Dr. Shufeldt *, who has given us a comprehensive sketch of
the anatomy of Heloderma, remarked that his own studies of the
Varanide convinced him “of the fact that Meloderma is far
removed from that group, having very little structural affinity
with it.” To these papers cited below, and to others quoted in
them, reference may be made for the views which have been
held with regard to the position occupied by the genus Heloderma
in the system.
In recently dissecting examples of these two genera I have
noticed two structural features in which the two genera are similar
and by which they may be differentiated from any other Lacer-
tilians whose anatomy is known, so far as concerns the points in
question. The first of these is a feature in the anatomy of the
respiratory organs which has indeed been described in Varanus
but not in Heloderma. As to the former genus, Meckel 7,
Giinther f, and Milani §, to whose investigations our knowledge
of the anatomy of the respiratory organs in the Varanide is
chiefly due, describe a short branch given off by the bronchus
shortly after it has entered the lung; this supplies the headward
extension of the lung which is so well marked in this genus of
Lizards. It is plainly figured by Milani||, whose illustrations 4,
particularly a diagrammatic figure, show that this twig arises im
front of an aperture in the walls of the intrapulmonary bronchus.
Some of Milani’s figures** also illustrate another somewhat im-
portant fact, which is that the bronchus until it gives off the branch
to which reference has been made does not really lie within the
the lung, but outside of it. The lung in growing forward has
wrapped round the end of the bronchus. Though apparently
within, this portion of the bronchus is really morphologically
outside of the lung. The interest attaching to the exact relation-
ship between the bronchus, the lung-tissue, and the first branch
of the bronchus appears to me to be this :—that this independent
branch arising so early from the bronchus 1s possibly to be compared
with the eparterial bronchus of the Mammalia. This comparison
is not suggested by Milani. In any case it is, so far as I am
aware, a structure that has not yet been described in any other
Lacertilian. A precisely comparable branch of the bronchus
occurs, however, in Heloderma, where its existence is an interesting
feature of resemblance to Varanus. I cannot find that any of
the writers TT who have described the lung of Heloderma have
noticed this—to my eyes, very striking—peculiarity of the lung.
Nor do the illustrations given by them show any signs of the
* P.Z.S. 1890, p. 233.
+ Deutsches Arch. f. d. Phys. 1818, Bd. iv. t P. Z. 8. 1861, p. 112.
§ Zool. Jahrb. (Abth. f. Anat.) vii. 1894, p. 581,
|| Zoe. cit. Taf. 31. figs. 16-18.
§| Loe. cit. p. 581, fig. R, Taf. 32. figs. 19-21.
** Especially fig. 20 of pl. 32 of his memoir.
++ Shufeldt, loc. cit. p. 202; Stewart, P. Z.S. 1891, p. 118; Miller, “The Structure
of the Lung,” Journ. Morph. vii. 1893, p. 170.
1907.] OF CERTAIN SPECIES OF SQUAMATA. 61
eparterial bronchus in Heloderma. ‘The length of the bronchi
and the complicated structure of the lungs themselves have,
however, been remarked upon, and in these matters Heloderma
also agrees with Varanus. The figure (text-fig. 16) shows the
commencement of the lungs in Heloderma suspectwm as seen
from the dorsal aspect. The long headward prolongation of
each lung, not perhaps sufficiently emphasised in Dr. Shufeldt’s
figure of these organs, is very reminiscent of Varanus. The
rings of the trachea are, as correctly stated by Shufeldt, in-
complete in the dorsal median line. The bronchi are described
by the same writer as “unusually long”; but it is not clear from
Text-fig. 16.
A portion of the trachea, the bronchi, and the upper parts of the lungs of
Heloderma, from the dorsal side.
The left lung cut open to show the course of the bronchus within it.
the description given, as Prof. Stewart * has pointed out, that this
great length does not apply to the intrapulmonary portion of
the bronchus. My own measurements of the length of the
bronchus up to the place at which it enters the lung agree with
those of Prof. Stewart in the case of Heloderma horridum. 1
find this length, in fact, to be as nearly as possible 13 mm. ‘To
be absolutely exact is impossible, on account of the pliability of
the bronchial rings and interspaces. The bronchi of Heloderma
are therefore shorter than those of Varanws (text-figs. 18 & 19,
* P.Z.S. 1891, p. 120.
62 MR. F, E. BEDDARD ON THE ANATOMY [Jan. 15,
pp. 65, 66), and not very much longer than those of a similarly-
sized Iguana tuberculata. Generally speaking, it is undoubtedly
correct to describe the Lacertilia as possessing short bronchi, to
which rule, indeed, Varanus offers the only very marked exception.
The relations of the bronchi to the lungs are not shown in
Shufeldt’s figure *, where the heart obscures the same, and are
wrongly shown in the figure of Millery. The latter author is
wrong (unless, indeed, the lungs of Heloderma suspectum, examined
by myself, are abnormal) in not indicating, in the figure referred to,
a conspicuous branch of the bronchus developed equally on both
sides of the body. When the trachea divides, the dorsal median
fibrous wall lying between the disjunct ends of the tracheal
semirings is continued down each bronchus. As Dr. Shufeldt has
remarked, the calibre of each bronchus is not far short of that of
the trachea itself. They are, in fact, particularly wide. The ends
of the bronchial semirings are, of course, visible on either side of
the median fibrous tract. ‘he bronchus approaches the lung and
becomes adherent to its mesial side and runs down in contact with
it for some distance until it finally enters the lung. At the point
of contact the upper ends of the’ semirings, 7. e. those lying
headwards, cease to be parallel with the lower ends and diverge
headwards. The dorsal membranous space ceases, and the semirings
in that section of the bronchus which is closely applied to the lung
embrace the lung. There is, in fact, a branching of the bronchus,
and this short branch may be seen to be lined by cartilaginous
semirings for a short distance into the interior of the lung. This is
not the case with the following apertures of communication between
the bronchus and the lung. I cannot but think that this branch
is comparable to that already referred to in Varanus. It 1s
further not without importance to notice that this “ eparterial
bronchus” in Heloderma is not serially comparable to the apertures
which place the cavity of the bronchus into communication with
the interior of the lung and which follow it. For the latter are
more ventral in position, as is plainly to be seen in the accompanying
figure (text-fig. 16). The ‘eparterial bronchus” is more dorsal
and is, in fact, lateral with reference to the main stem of the
bronchus.
The figure (text-fig. 16) which illustrates the branching of the
bronchus before entering the lungs also shows on the left side
the interior of the lung as seen when the bronchus is slit up after
it has given off the branch referred to. I have thought it worth
while to introduce this view of the lung of Heloderma, since the
figure given by Miller ¢ does not appear to me to represent quite
accurately the mode of communication between the interior of
the bronchus and the lung-substance, nor does he indicate the
adherence of the bronchus to the lung for a considerable distance
before entrance. He does, however, illustrate the important fact
* Toe. cit. pl. xvi. fig. 3. + Journ. Morph. 1898, pl. vil. fig. 5.
t Loe. cit. pl. vii. fig. 5.
1907.] OF CERTAIN SPECIES OF SQUAMATA. 63
that the bronchus traverses a considerable distance within the
lung before it disappears. My own illustration will show that
the semirings of the bronchus are complete for a considerable
distance, and perfectly easily recognisable, since they show no
particular differences from the semirings in the extrapulmonary
region of the bronchus. The bronchus communicates with the
lung by copious apertures, which are not situated in the region
of the bronchus corresponding to the fibrous band which unites
the tips of the semirings in the extrapulmonary region of the
lung; these apertures would seem to be rather breaks in
continuity of the semirings themselves. Their disposition is
thus reminiscent of the way in which the rudimentary lung of
certain Snakes arises from the bronchus. There is a simple hole
in the bronchus which leads into the lung in the case of those
Snakes.
In a paper communicated to this Society a good many years
since * J described the complicated branching of the cystic duct
and its anastomosis with the hepatic duct in Varanus salvator.
T have since then discovered that the same network, comparable
to that which is found so generally among the Ophidia, occurs
also in V. gould, though it is in that species rather less developed
than in V. salvator. Quite recently I have dissected out the bile-
ducts in V. nloticws, of which species I have had the opportunity
of examining several very small examples preserved in spirit.
Thad one of these injected from the gall-bladder, and the injection
(chrome-yellow rubbed up im olive-oil) ran readily along the
branches of the cystic and hepatic ducts. The accompanying figure
(text-fig. 17, p. 64) is fairly accurate (but I fear not absolutely
so) as regards the network, which, as will be seen, is much like
that of V. salvator, but perhaps not quite so complicated. More-
over, when once the hepatic and cystic ducts have left the surface
of the gall-bladder there are apparently no further anastomoses
between them, as there are—though to a limited extent—in
V. salvator. On the other hand, there are some species of Varanus
in which there is no network formed by the bile-ducts on their
emergence from the gall-bladder (as is also found among the
Ophidia). To some of these I have referred in my communication
just quoted. I have since carefully examined Varanus exanthe-
maticus, and find that the cystic duct emerges as, and continues
to be, a simple duct throughout. ‘The same is the case with
V. griseus. It is not wise perhaps to generalise on these few
data; but so far as the facts go they agree with an important
external character by which the species referred to may be
grouped. In JV. salvator, V. gouldit, and V. niloticus the nostril is
a circular aperture, while in the other species mentioned it is
obliquely placed and slit-like.
In all the species of Varanus that have heen referred to in the
* P.Z.S. 1888, p. 106. The illustrative figure (fig. 4, p. 105) has been copied in
Gegenbaur’s ‘ Verg]. Anat. Wirbelth.’
64 MR. F. E. BEDDARD ON THE ANATOMY [Jan. 15,
present communication the pancreas is a solid body, as shown in
the figure of that of V. niloticus (text-fig. 17). In all of them a
Text-fig. 17.
~
Pancreas, gall-bladder, &c. of Varanus niloticus, to show the course of
the bile-ducts. :
g-b. Gall-bladder; 4.d. Hepatic ducts; Z. Liver; O. Orifice of conjoined bile-ducts ;
P. Pancreas; P.v. Portal vein.
very slender splenic lobe arises near to the anterior end of the
pancreas and passes to the spleen*. The species, however, show
* In Varanus exanthematicus the end of the splenic not merely touches, but
enwraps and is enwrapped by the spleen. This intimate relation between the
spleen and the pancreas recalls a similar close association occasionally found among
the Ophidia.
OOK OF CERTAIN SPECIES OF SQUAMATA. 65
certain differences among themselves in the position of the
duodenum with reference to the pancreas. In J. niloticus the
pancreas is separated by a considerable tract of mesentery from
the duodenum, which results in the exposure of a long pancreatic
duct. The same is the case with V. ocellatus. On the other hand,
in V. griseus and V. exanthematicus the end of the pancreas
touches the duodenum close to the point of entrance of the
pancreatic and bile ducts, and there is therefore no great length of
pancreatic duct.
Text-fig. 18.
Lungs of Varanus exanthematicus.
Bronchi opened to show exit of “eparterial bronchi” close to pulmonary
artery (P.a.).
In his figures of the lung of several species of Varanus, Milani
has pointed out certain differences which distinguish them. Thus
the lungs of V. bengalensis are lobed externally in a fashion which
is not to be found in the other species which are described. In
examining, for the purposes of the present communication, the
lungs of several species of Varanus, I have observed two small .
points of difference between the lungs of certain species which are
not referred to by Milani. Of these both involve an asymmetry
Proc. Zoou. Soc.—1907, No. V. 5
66 MR. F. E. BEDDARD ON THE ANATOMY [Jan. 15,
or symmetry of the lungs and the windpipe as the case may be,
not, indeed, in point of length, which is a common feature among
Lizards and one of the most obvious features in which they
approach the Snakes. This asymmetry, when it occurs, affects
the position of the “ eparterial bronchus” and the ventral forward
projection of the lung between the bronchi. In Varanus griseus
(text- fig. 19) the left lung, as well as the right lung, sends forward
towards the bifurcation of the bronchi a thin diverticulum, which
has an entire cavity not divided by any meshwork and seems to
be comparatively unvascular. ‘This can readily be lifted up and is
seen to be not attached to the bronchus of the lung of which it
is an outgrowth.
Text-fig. 19.
Lungs of Varanus griseus. Details as in text-fig. 18.
So far, therefore, the lungs are symmetrical. But the origin of
the eparterial bronchus is not symmetrical. On the right side
this branch arises from the bronchus at a distance of 27 mm. from
the point of bifurcation of the bronchi; in the case of the left
90 OF CERTAIN SPECIES OF SQUAMATA. 67
lung this distance was only 21 mm. <A second specimen showed
precisely the same relations in all these points of structure.
Contrasted with this (compare text-figs. 18, 19) are the different
conditions observable in Varanus exanthematicus. In the latter
species (text-fig. 18, p. 65) the two branches of the bronchi were
exactly symmetrical and each was situate 33 mm. from the bifurca-
tion of the bronchi. Only the right lung gave off a forwardly
directed lobe situated on the inner side of its bronchus. There was
nothing to correspond in the left lung. I have not been able to
compare these conditions with those of many other species of
Varanus. But in both JV. ocellatus and V. niloticus there was
precisely the same asymmetry in the relative positions of the
branch of the bronchus, which in all cases lies behind the aorta of
its side.
Summary of more important new Facts contained in this
Communication.
In view of the fact that very few genera and species of
Lacertilia have been studied anatomically, it is a little difficult at
present to differentiate between more and less important structural
details as evidence of affinities between different genera. The
following réswmé, therefore, will be necessarily only an attempt to
lay stress upon what appear at present to be the more important
new facts which I have set forth in this communication.
(1) The pancreas in the Lacertilia, as already known, differs in
different genera. I have added to the existing knowledge some
new facts with regard to genera and species not examined by
others. It appears from this that the chief variability in the
ancreas consists in the presence or absence of a splenic lobe and
in the relations of the latter to the spleen. The classificatory
importance of the facts does not appear to be great ; since, though
the Iguanoids, Zgwana and Liolemus, are like each other in the
relations between the splenic lobe of the pancreas and the spleen,
we find in Varanus and Chameleon differences between different
species in these points. The pancreas is nearly always a compact
gland; but not so in Zonurus.
2) The variations in the structure of the viscera among the
Chameleons concern principally the proportions between the two
lobes of the liver, the form of the diverticula of the lungs and the
absence or presence of these, the degree of pigmentation of
the body-cavity, and the degree of coiling of the intestines.
(3) The variations in the structure of the viscera in the
different species of Varanus concern principally the presence or
absence of a bile-duct network and certain minute differences in
the lungs. It seems possible that those species with a round
nostril are distinguished from those with an oblique slit-like
nostril by the possession of this network.
(4) The simplicity of structure which is often associated with
small-sized forms as compared with their allies of larger size is
re
a"
68 MR. G. H. KENRICK ON PYRALIDE [Jan. 15,
well seen in the two small species of Chameleons, viz. Ch. pumilus
and Ch. teniobronchus, where the lungs have no diverticula and
the intestinal tract is nearly straight.
(5) The very general presence of a gubernaculum cordis among
the Lacertilia renders its absence in Zonurus a matter worthy of
comment.
(6) The most important fact, perhaps, which I have been able
to ascertain is the persistence in several genera of Lacertilia
of considerable remains of the fourth visceral arch (second
branchial). This is a fair-sized bar of cartilage which does not
make any connection with the copula below. The existence of
this arch has, however, been recorded in the adult Lacerta by the
late W. K. Parker.
(7) The two genera Varanus and Heloderma (which are quite
remote from each other in some structural features) agree with
each other in that each bronchus is adherent to its lung for
some little distance before it enters it, and emits a short branch to
the upper end of the lung before it becomes confluent with
the lung.
(8) It is interesting to note the double gastro - hepatic
membrane in certain Scincide, which is associated with a
correspondingly double set of gastro-hepatic veins, as distinctive
of that family, though not universal.
4, A List of Moths of the Family Pyrahde collected by
A. E. Pratt in British New Guinea in 1902-3, with
Descriptions of new Species. By Grorce H. Kenrick,
F.Z.S.
[Received December 8, 1906. |
(Plates III. & IV.*)
This collection was made under circumstances mentioned by
Mr. Pratt in his book ‘ Two Years among New Guinea Cannibals,’
published in 1905, and beyond the fact that most of the speci-
mens were taken at light very little information can be given.
The country in which the collections were made appears in
some of its characters to resemble Darjiling: there are the same
precipitous ridges with narrow valleys between, all with a back-
ground of snowy mountains of great elevation, and everywhere
there is much dense forest. The climate, with its abundant rain
in the wet season and brisk air in the dry season, is also similar,
while an abundant lepidopterous fauna completes the resemblance.
Although most of the insects were taken at light, in most cases
females were well represented and the condition of the insects is
extremely good.
* Hor explanation of the Plates, see p. 87.
12 BS, USO, IPA. ih,
36
EC. Knight delet lith, West,Newman chromo.
PYRALIDA, FROM BRITISH NEW GUINEA.
124, SAO, IL INV.
~ om SR a ae em Vers ta
Sem i (0 Wn me) em
West, Newman chromo.
E.C. Knight del.et Ii
A.
Steal
5
se
GUIN.
NEW
PROM BRITISH
PYRALIDA:
1907. | FROM BRITISH NEW GUINEA. 69
Subfam. SCH@NOBIIN.
1. CIRRHOCHRISTA STHERIALIS Led.
2. C. PULCHELLALIS Led.
3. C. FIGURATALIS WIk.
4, C. CACONALIS Swinh.
MonopontTA, gen. n.
Palpi porrect, more than length of head, parallel, clothed with
hairs, but very thin; antenne simple; patagia shorter than thorax.
Venation of fore wing: veins 2 and 3 before end of cell, 4 and 5
from end of cell, 6 and 7 from end of cell, curved and nearly
parallel, 9 and 10 from cell. Hind wing: cell rather short ; veins
2 and 3 before end of cell, 4 and 5 from end of cell, 6 from upper
angle, 7 anastomosing with 8 halfway between end of cell and
end of wing; costa slightly curved; hind margin convex, rounded
at angle.
Type, J. passalis.
5. MoNnoponta PASSALIS, sp. n. (Plate IV.)
Head, legs, thorax, and patagia dark brown, abdomen rather
paler. Fore wing pale brown inclined to purplish, an oblique
dark line at one-fifth of the length of the wing, followed by a
much darker portion outwardly bounded by a curved dark line
deeply notched near angle of wing, this is followed by a pale
ochreous shade. Hind wing pale ochreous at base, shading into
darker towards the margin and having an indented dark line
similar to that of the fore wing; fringes darker.
Exp. 50 mm.
Hab. Kebea, one specimen.
Subfam. PHycrrin 2.
6. PHycrra pinawa, sp. n. (Plate IIT.)
Head, legs, palpi, and body ochreous, darker above. Fore
wing marbled with purple, green and gold, the raised scales
beyond the cell darker; a curved postmedian line dark margined
with paler. Hind wing plain purplish grey. Fringes of fore wing
purple ; fringes of hind wing paler.
Exp. 24 mm.
Hab. Dinawa, four specimens.
7. EYIELLA FUSCALIS, sp. n. (Plate ITT.)
Head, legs, palpi, and body brown with numerous pinkish hairs.
Fore wing pinkish brown, a darker brown mark two-thirds along
the inner margin edged on either side with paler; fringes dull
pink. Hind wing smoky grey; fringes same colour.
Exp. 28 mm.
Hab. Dinawa and Kebea, four specimens.
8. HypstpynA roBusta Moore.
70 MR. G. H. KENRICK ON PYRALIDE [Jan. 15
Subfam. EHPrPaASCHIINé.
9. Macauua @NnocHRoa Hmpsn.
10. M. prcta Warr.
11. M. nycricHroaLis Hmpsn.
12. M. semMINIVEA W1k.
13. M. marearira Butl.
14, M. UNIPUNCTALIS, sp. n. (Plate IIL.)
Head, legs, palpi, and antennze ochreous, tarsi barred with
darker. Fore wing reddish ochreous, with wavy antemedian and
postmedian lines dark brown, between them is a black spot very
near costa; the outer third of the wing is brown and there is a
marginal row of black dots. Hind wing much paler, with similar
marginal row of black dots and a faint curved median line.
Fringes ample, shining brown.
Exp. 28 mm.
Hab. Dinawa, one specimen.
15. M. CARADRINIFORMIS, sp. n. (Plate ITT.)
Head, antenne, palpi, and legs ochreous, tarsi darker; thorax,
patagia, and fringes all of the same colour. Fore wing ochreous,
with curved and rounded postmedian line paler but outwardly
margined with darker ; on the costa are three equidistant badly
defined, slightly darker markings; from the middle of the base
of the wing is a ragged darker streak, and this becomes stronger
as it approaches the postmedian line near the end of which is
a dagger-like marking. Hind wing uniformly ochreous.
Exp. 32 mm.
Hab. Dinawa and Kkeikei.
16. M. pomauis, sp.n. (Plate III.)
Head, thorax, palpi, and processes pale green, tibie barred
with pale green and brown; eyes black with a network of buff ;
abdomen green above, buff below. Fore wing apple-green with
irregular markings of dark chestnut; a transverse band at one-
third length of wing, an irregular divided blotch on inner margin
reaching nearly to the angle, and a square-shaped blotch on costa
near, but not at, the tip of the wing; a dark spot at end of cell.
Hind wing pale brown ; fringes paler.
Exp. 24-26 mm.
Hab. Dinawa.
17. M. cuRTULALIS, sp. n. (Plate ITT.)
Head, antenne, palpi, and legs pinkish buff, tarsi ringed with
darker. Fore wing wainscot-colour with a conspicuous chocolate
1907.| FROM BRITISH NEW GUINEA. 71
tip; three dots on costa before this mark and a dot at end of
cell. Hind wing pale clouded on margin with smoky.
Exp. 23-26 mm.
Hab. Dinawa.
18. M. apicauis, sp. n. (Plate IIT.)
Head, antennee, and palpi chestnut, thorax and abdomen the
same with a few grey hairs; patagia paler. Fore wing; ground-
colour chestnut, but the band, which is conspicuously white, is
extended in some specimens in a tooth into the hind margin while
on the other side it produces a basal blotch. Hind wing fuscous
edged with darker. Fringes of both wings pale chestnut.
Exp. 30-34 mm.
Hab. Kebea, Dinawa, Babouni, Ekeikei.
19. M. TENEBROSALIS, sp. n.
Head, legs, palpi, and antennz ochreous; abdomen grey, with
dark transverse band on second segment. Fore wing ochreous with
basal and apical blotch; the band begins near the base with a dark
interrupted line, it is twice as wide on costa as on inner margin
and the exterior boundary is an oblique angulated dark line; the
interior of the band is reddish ochreous with a dark dot at end
of cell. Hind wing smoky. Fringes of both wings ochreous.
Exp. 32-36 mm.
Hab. Kebea, Ekeikei, Babouni.
20. M. PERDENTALIS, sp. n. (Plate III.)
Head, legs, palpi, and antenne pinkish ; thorax, patagia, and
abdomen ochreous grey, with a slight chestnut band on second
segment. Fore wing ochreous grey with dark lines and chestnut
patches, basal blotch grey ; band very irregular, bounded on
inner side by an oblique dark angulated line, on outer side by a
strongly marked line, very narrow on inner margin but occupying
about one-third of costa and mostly chestnut; beyond the band
is a darker cloud towards apex ; there is also a dark central dot.
Hing wing smoky with darker veins, rather paler towards the
base. Fringes of both wings pale grey.
Exp. 30 mm.
Hab. Dinawa, Kebea.
21. M. PORPHYREALIS, sp. n. (Plate IIL.)
Head, legs, palpi, and antennz pinkish ochreous ; thorax grey
to reddish. Fore wing mostly dark brown, with whitish and
grey interrupted transverse bands; the median band is very
indistinct but its outer boundary is marked by a paler narrow
angulated band with a tooth in the middle; there is a row of
subterminal dark spots. Hind wing straw-colour with darker
margin. Fringes of both wings pale.
Exp. 34-36 mm.
Hab. Ekeikei, Kebea, Dinawa.
72 MR. G. H. KENRICK ON PYRALIDE [Jan. 15,
All the species of Macalla enumerated above exhibit considerable
colour variation and in most cases a certain amount of sexual varia-
tionas well. It is quite possible that when they are better known
some may prove to be varieties; on the other hand, it is also
possible that some now treated as varieties may turn out to be
veritable species.
22. PoLYLOPHOTA BARBAROSSA Hmpsn.
23. P. TRUNCALIS, sp.n. (Plate IIT.)
Top of head, antennz, palpi, legs, and thorax pale buff with
some black hairs; patagia black. Fore wing buff with a darker
cloud near the angle and an interrupted subterminal double line;
a subtriangular dark mark extends from near the base along the
costa gradually thinning out; in the midst of this is the transverse
brush of raised scales nearly black. Hind wing reddish with
dense rufous hairs near body. Fringe of fore wing very dark ;
fringe of hind wing reddish. Tibiz of hind legs clothed with
rufous hairs.
Exp. 38 mm.
Hab, Dinawa.
24, LOcASTRA CASTANEALIS, sp.n. (Plate IIT.)
Head, legs, antenne, and palpi pale chestnut ; tarsi ringed
with darker ; thorax and patagia chestnut, abdomen paler. Fore
wing chestnut, the first third suffused with grey near the inner
margin, after this comes a dark transverse angulated line forming
the inner boundary of the band, the outer boundary of which is
a curved pale transverse line; at the angle of the wing is a
conspicuous grey elliptical blotch. Hind wing grey; fringes paler.
Exp. 36 mm.
Hab. Dinawa.
25. STERICTA SPORETA Turn.
26. S. HARRALDUSALIS W1lk.
27. S. prRAsSINA Warr.
28. S. DIvITALIS Guen.
29. S. FLAMMEALIS, sp.n. (Plate IIT.)
Head and palpi greenish, antennze and legs pink, tarsi barred
with darker, thorax and process greenish; body pinkish. Fore
wing: ground-colour dark olive-brown; at one-third distance
along the costa is a broad transverse band and at one half a
distinct angulated but narrower band; these markings and a
portion of the wing near the hind margin are of pale olive-green.
Hind wing rich golden orange. Fringes of both wings orange-
pink.
Exp. 26 mm.
Hab. Ekeikei, one specimen.
1907.] FROM BRITISH NEW GUINEA. 73
30. S. cornucaLis, sp. n. (Plate IIT.)
Head, palpi, antenne, legs, and thorax rufous buff. Fore wing
buff with darker and lighter lines and dots; on the costa a dark
mark at base and another at one-third length of wing, smaller
dots at one half and two-thirds, a cloud at the tip; a wavy
subterminal line dark edged with lighter; a dark dot on inner
margin near body. Hind wing buff, with a dark wavy line edged
with lighter and dark central spot; fringes buff spotted with
darker.
Exp. 22 mm.
Hab. Dinawa.
31. S. SUBVIRIDALIS, sp. n. (Plate ITI.)
Head, legs, thorax, abdomen, palpi, and the long processes
which extend to the back of the thorax, buff. Fore wing dull
brown, with two transverse pale greenish bands parallel to hind
margin, the first at one-half and the second at two-thirds length
of wing; below the cell these bands run into a large patch of the
same colour which continues along two-thirds of inner margin
and projects a little beyond the second transverse band. Hind
wing dull brown ; fringes same colour.
Exp. 28 mm.
Hab. Kebea, Dinawa, Babouni.
32. ORTHAGA VITIALIS Wlk.
33. O. POLYCHROALIS, sp. n. (Plate ITT.)
Head, antenne, legs and thorax chestnut inclining to pink ;
abdomen rather paler, with no band on second segment. Fore wing
greenish with chestnut patches; basal blotch mingled olive and
chestnut, with raised scales in longitudinal dark line, then a vertical
line edged outwardly with white; the band is badly defined but
there is an angulated thin line forming the external boundary ;
the remainder of the wing is greenish white, with a chestnut
apical patch. Hind wing pale buff. Fringes of both wings pale ;
underside of costa of both wings broadly red.
Exp. 30 mm.
Hab. Dinawa.
34. O, CHIONALIS, sp.n. (Plate III.)
Palpi simple in both sexes, reaching vertex of head ; antennz of
male ciliated and with a process consisting of an immense tuft of
scales reaching over part of thorax; in the female this is very
small.
Head, palpi, antenne, and legs pinkish buff, legs ringed with paler
at the joints; thorax, patagia, and process greenish; abdomen
greyish ochreous. Fore wing: ground-colour dark olive-brown,
basal patch with raised scales both light and dark followed by an
oblique narrow white band, then follows the main band, in some
74 MR. G. H. KENRICK ON PYRALIDE [Jan. 15,
specimens suffused with white and bounded by a curved paler
line after which is a darker cloud. Hind wing fuscous; fringes
of both wings paler.
Exp. 38-39 mm.
Hab. Ekeikei, Babouni, Kebea, Dinawa.
39. ORTHAGA FUSCOFASCIALIS, sp. n. (Plate IIT.)
Antenne of male ciliated, with process roughly scaled and
reaching thorax ; female without process.
Head, palpi, antenne, legs, and top of thorax ochreous; the
male has a dark collar to thorax, absent in the female ; abdomen
ochreous. Fore wing: ground-colour fuscous; a dark basal blotch
occupies one-third of the wing; the band is bounded on the
inner side by a narrow vertical white band, this is followed by
pale ochreous with dark dots on the costa and is bounded out-
wardly by a dark wavy interrupted line; the remainder of wing
of the ground-colour with a row of subterminal triangular dots.
Hind wind fuscous ; fringes paler. In some specimens the band
is suffused with chestnut.
Exp. 26-30 mm.
Hab. Dinawa and Kebea.
36. O. COLUMBALIS, sp. n. (Plate ITI.)
Head, legs, antennze, thorax, and body dove-colour ; legs ringed
with black. Fore wing dove-colour, on costa at base a triangular
black patch, the apex of which reaches the inner margin as a
thick black line and forms the inside vertical boundary of the
band; the band itself is dove-colour, but there is a thin curved
black line parallel with inside boundary and an irregular black
patch on vein 2; the outer boundary of the band is a curved
thick black line, and beyond this on the costa is an anteapical
black patch. Hind wing dove-colour ; fringes paler. Underside
smoky with whitish dots on costa.
Distinct from seminivea in absence of white thorax and black
collar.
Exp. 36 mm.
Hab. Kebea, one specimen.
37. TERMIOPTYCHA CYANOPA Meyr.
Subfam. ENDOTRICHINE.
38. ENDOTRICHA ALBICILIA Hmpsn.
39. KE. persicopa Meyr.
40. CovACHENA TRIVITRALIS Hmpsn.
41. C. atuensis Butl.
~I
OX
1907.] FROM BRITISH NEW GUINEA.
TIPULIFORMA, gen. n.
Fore wing: vein 2 present; veins 3, 4,5 from lower end of
cell; veins 6, 7 from upper end of cell; vein 7 parallel and close
to 8 for one-third of its length, then turning down; vein 9 from
middle of cell. Hind wing: veins 7 and 8anastomosing. Proboscis
well developed ; palpi upturned reaching vertex of head, densely
clothed with hairs ; antennz simple, three-quarters of the length
of fore wing. Abdomen projecting for half its length beyond
hind wing; legs long and slender, with small spurs on the tibie ;
fore tarsi somewhat hairy. Fore wing long and narrow; hind
wing subquadrate.
42. 'T. TRIANGULALIS, sp. n. (Plate III.)
Thorax, head, and palpi grey ; patagia whitish ; underside
quite white; abdomen grey above, white beneath, tuft white;
eyes very prominent. Fore wing dark smoky-grey shot with
reddish; costa narrowly buff for two-thirds of wing; a small basal
spot white, followed by three other white dots along the costa ;
the inner margin with a narrow pale streak expanding into a
quadrangular blotch at the angle. Hind wing white; a marginal
dark band tapering from apex of wing and interrupted by a tooth
in the middle. Fringes of both wings dark at apex and white at
angle.
Exp. 36 mm.
Hab. Dinawa.
Subfam. PyRaLin#®.
43, VITESSA SURADEVA Moore.
44, V. zemirA Cram.
45. V. GRISEATA, Sp. n. (Plate IIT.)
Head, thorax, and patagia pale orange, with four black dots
across the front of thorax; abdomen black ringed with yellow;
palpi black. Fore wing pale greenish grey ; veins near the hind
margin tinged with black; a vertical metallic blue streak across
the base of the wing and the space between this and the body
dull orange, with one blue spot and a blue oblique streak below
it; fringes same colour as fore wing. Hind wing black; fringes
paler.
Exp. ¢ 51 mm., 2 66 mm.
Hab. Dinawa.
46. HypsopyGia postruAvA Himpsn.
47, SACADA INORDINATA W1k.
48. S. nigRopuncTata Hmpsn.
76 MR. G. H. KENRICK ON PYRALIDE [Jan. 15,
Subfam. HypDRocAMPIN«A.
49, CyMoORIZA USTALIS WIk.
50. AULACODES PLICATALIS W1k.
51. A. GONIOPHORALIS Hmpsn.
52. A. ACROPERALIS Hmpsn.
53. A. DIOPSALIS Hmpsn.
54, A. TRICHOCERALIS Hmpsn.
55. A. LUNALIS, sp.n. (Plate III.)
Head, thorax, and patagia buff; first segment of abdomen brown
followed by a pale band, the remainder golden. Fore wing dark
brown ; a pale brown stripe runs from the body and reaches the
costa about midway; it is then continued as a fine line along
the costa to the apex; along the inner margin is a whitish
stripe with an angulated upward projection towards the middle
of the wing; on the disc is a crescent-shaped white mark and
this is followed by a white subterminal line with an orange band
beyond ; fringes pale. Hind wing: the base for about one-third
of the wing is white, the remainder orange; on the margin are
four black dots, the two exterior having white spots above them ;
in the middle of the wing are two black lines nearly parallel to
hind margin.
Exp. 20 mm.
Hab. Dinawa.
56. A. PURPUREALIS, sp. n. (Plate III.)
Head, thorax, and abdomen purplish grey ; fore legs pale ringed
with dark. Fore wing purplish grey with orange markings; a
curved irregular band beginning on the inner margin and joining
an orange band along the hind margin which is preceded by a
silvery band; nearer the base is a dusky orange patch followed
by a crescent of purple and then a long triangular mark of orange.
Hind wing purplish grey at base, with a broad median band
followed by a darker band, and finally by a narrow orange marginal
band in the middle of which are seven silver-studded black spots ;
fringes paler.
Exp. 25 mm.
Hab. Dinawa.
57. A. BIPUNCTALIS, sp. n. (Plate IIT.)
Head pale brown, thorax and first segment of abdomen dark
brown, followed by a slightly paler band; legs pale ringed with
dark brown. Fore wing dark brown; with a white streak fading
into buff on the inner margin ; a white subterminal line followed
by an orange band; fringes yellow. Hind wing: basal third
white, remainder orange, with a short white band bordered with
dark near the angle and a dark line parallel to the first but nearer
1907. ] FROM BRITISH NEW GUINEA. Ct
to the base of wing; three black dots on the hind margin, two of
which have white dots above them.
Exp. 23 mm.
Hab, Dinawa.
58. PARTHENODES RECTANGULALIS, sp. n. (Plate IV.)
Head, legs, palpi, and patagia white; thorax black above;
antenne and abdomen dark brown. Fore wing: costa straight,
hind margin nearly at right angles for half the width of wing,
then slightly oblique to angle of wing; ground-colour white, with
black border to hind margin and a parallel transverse narrow
band at one-third of wing; from where this meets the inner
margin is a narrow oblique red band to base of costa; from the
lower end of black line on the hind margin is a double red oblique
streak reaching costa and extending to middle of wing, becoming
smoky on reaching the inner margin. Hind wing: base and hind
margin white, followed by a smoky band, becoming red nearer
the body and containing three white specks, slightly angulated.
Fringes white.
A very striking species both in form and colour.
Exp. 24-30 mm.
Hab. Kkeikei, Dinawa.
59, TALANGA TOLUMNIALIS WIk.
60. BRADINA IMPRESALIS Led.
61. B. MELANOPERAS Hmpsn.
62. B. GuaucinaAuis Himpsn.
63. ORPHNOPHANES EUCERALIS WIk.
64. CoProBASIS MESOPSECTRALIS Hmpsn.
65. DRACENURA ALBONIGRALIS Hmpsn.
66. D. HoRocHROA Meyr.
67. D. ToRRIDALIS, sp. n. (Plate IIT.)
Head and thorax fuscous; abdomen and posterior half of
patagia pale buff, last three segments of abdomen fuscous; tuft
buff; legs and underside white. Fore wing fuscous shot with
purple, a dark central spot and a transverse dark mark at end of
cell; beyond this is a curved thin dark line; fringes paler. Hind
wing yellowish, the extreme hind margin darker, shading
gradually into the colour of the wing and almost destitute of
fringe in the male.
Exp. 33 mm.
Hab. Kebea.
68. D. UNICOLORALIS, sp.n. (Plate IV.)
Vertex of head pale, palpi and thorax dark grey; posterior
end of patagia and abdomen pale; tuft black and a thin black
78 MR. G. H. KENRICK ON PYRALIDEH [Jan. 15,
lateral fringe. Fore wing olive-grey shot with purple, a minute
black apical dot and fine black dots on hind margin; fringe pale.
Hind wing: outer margin as fore wing; internal area straw-
colour.
Exp. 36 mm.
Hab. Dinawa, one specimen.
69, PILETOCERA INCONSPICUALIS, sp. n. (Plate IV.)
Fore tarsi naked.
Head, legs, antenne, tarsi, thorax, and abdomen dark brown ;
tuft whitish orange. Fore wing dark brown with faint purplish
tinge ; an antemedian, median, and postmedian angulated trans-
verse line darker; a white dot at end of cell and another in the
middle of cell, also a faint dot where the postmedian line reaches
the costa. Hind wing with two angulated darker lines. Fringes
dark with lighter patch at angle of wing.
Exp. 30 mm.
Hab. Kebea.
Subfam. PyRAUSTIN®.
70. PYCNARMON JAGUARALIS Guen.
71. P. vactiFERALIS W1k.
2, EUTEPHRIA CRIBRATA Fabr.
3, ZINCKENIA FASCIALIS Cram.
74, HETrEROCNEPHES SCAPULALIS Led.
75. REHIMENA PHRYNEALIS WIk.
76. AGROTERA PICTALIS Warr.
77. A. FuMosA Himpsn.
78. A. EUDOXANTHA Hmpsn.
79. A. FLAVIBASALIS Hmpsn.
80. A. SEMIPICTALIS, sp. n. (Plate IV.)
Head, thorax, and front of abdomen yellow spotted with orange,
last three segments of abdomen purple; tuft orange. Fore wing:
basal portion pale yellow flecked with orange, abruptly and
obliquely terminated by a patch of rich brown shot with purple
occupying the rest of the wing, in the middle of which is a red
irregular spot ; the costa towards the apex is paler, and there is
an obscure darker curved line in the middle of the wing. Hind
wing similar to the fore wing, but without the red spot. Fringes
conspicuously white with dark interruptions, giving the idea of a
scalloped margin.
Exp. 30 mm.
fab. Dinawa.
81. PagyDA PARAPHRAGMA Meyr.
1907. ] FROM BRITISH NEW GUINEA. 79
82. CNAPHALOCROCIS MEDINALIS Guen.
83. MARASMIA FUSIFASCIALIS Hmpsn.
84, RHIMPHALEA HERANIALIS WI1k,
85. R. sceLatTatis Led.
86. R. LINEALIS, sp. n. (Plate IV.)
Head and thorax golden; palpi, antenne, and legs brown ;
patagia darker; abdomen golden, but darker on the margin of
each segment; tuft golden. Fore wing whitish with purple
gloss and margins edged with gold, with numerous connected
streaks and blotches of purplish brown. Hind wing similar, but
the chief blotch is at the apex and a secondary blotch at angle.
Fringes paler.
Exp. 50 mm.
Hab. Dinawa.
87. SYNGAMIA FLORIDALIS Zell.
88. S. AMPLIATALIS Led.
89. S. MARMORATA Lucas.
90. BoccHORIS TELPHUSALIS WI1k.
91. B. aprpawis Led.
92. B. INVERTALIS Snell.
93. B. AuRoTINCTALIS Hmpsn.
94. B. EUPHRANORALIS W1]k.
95. PrmocRocIs ANIGRUSALIS W1k.
96. P. copropasis Hmpsn.
97. ULOPEZA CRUCIFERALIS, sp. n. (Plate IV.)
Head, antennee, legs, and palpi bright ochreous; abdomen same
colour in female, but lighter im male; tuft pale. Fore wing
ochreous clouded with purple; three equidistant spots on eosta
are produced in widening angulated bands until they meet a
longitudinal band on inner margin; in the middle of the outer
band is a projection reaching the hind margin. Hind wing
yellow ; fringes same colour.
Exp. 26 mm.
Hab. Ekeikei, Mafalu.
98. NosopPHORA DISPILALIS Himpsn.
99. N. FuLvALIS Hmpsn.
100. N. semrrrirauts Led.
101. N. BARBATA Hmpsn.
102. CHALCIDOPTERA EMISSALIS W1k,
80 MR. G. H. KENRICK ON PYRALIDE [Jan. 15,
103. CAPRINIA FELDERI Led.
104. C. congropaTaLis W1k.
105. C. cAsTANEALIS, sp. n. (Plate IV.)
Head, thorax, and abdomen above dark grey, a white band on
first segment of abdomen. Fore wing black at base followed bya
white band for one-third of wing, this is edged with black and the
tip of the wing is also black, between them the space is bright
chestnut ; on the costa and extending to bottom of cell is a white
spot and there is a white dot on the inner margin. Hind wing:
basal half white, remainder chestnut; margin broadly darker.
Fringes dark.
Exp. 34 mm.
Hab. Ekeikei, Kebea, Mafalu.
106. C. oceLLauis, sp. n. (Plate IV.)
This insect resembles Rhimphalea trogusalis W1k., but differs in
the following particulars :—The spot in the cell is developed into
a distinct ocellus with white centre, black ring, and yellow space
with outer dark line, and it is much more vivid on the under-
side; the postmedian line is double with a yellow centre, and
there is a distinct parallel subterminal line, the hind margin
being yellowish with a dark cloud in the centre.
Sir G. Hampson considers that its right place is in the genus
Caprinia.
Exp. 27 mm.
Hab. Ekeikei.
107. FrnopEs XANTHALIS Hmpsn.
108. TygPANODES RADIATA, sp. n. (Plate ITT.)
Head, thorax, and tuft orange ; collar black; legs grey. Fore
wing grey with whitish hyaline patches and streaks; a dark spot
at base of costa followed by pale orange, then a transverse narrow
black band, after this a triangular white patch from costa to
inner margin, the nervures 1 and 2 showing on this; beyond is a
large irregular white patch, the apex being broadly dark grey ;
fringe greyish yellow. Hind wing dark with the nervures
whitish and an oval white patch beyond the middle; fringe
greyer than in the fore wing. ‘
Exp. 44 mm.
Hab. Kebea.
109. NEvVRINA PROcoPIA Cram.
110, PHRYGANODES MACULICOSTALIS Hmpsn.
111. P. euypHopatis Wk.
112. P. opscuratTa Moore.
113. P. Basauticatis Led.
114. P. oponrosticra Hmpsn.
1907. | FROM BRITISH NEW GUINEA, 81
115. P. mMarcarira Butl.
116. DicHocrocis xANTHOCYMA Hmpsn.
117. D. Bvaxatis W1k.
118. D. pUNCTIFERALIS Guen.
119. D. NiGRILINEALIS WIk.
120. D. PARDALIS, sp. n. (Plate IV.)
Head and palpi orange; legs paler; tarsi with dark rings ;
antenne dark; thorax ochreous, spotted with dark brown ;
abdomen ochreous with darker cloud on back; tuft dark. Fore
wing pale ochreous, with narrow dark basal band and two narrow
equidistant bands crossing from costa to inner margin where they
are wider or branched; a subterminal dark patch which only
extends across half the wing. Hind wing ochreous slightly shot
with purple, a dark subterminal line and a narrow dark trans-
verse line across half the wing. Fringes golden. All the markings
variable.
Exp. 30-40 mm.
Hiab. Mafalu.
121. D. BrmacuLatis, sp. n. (Plate IV.)
Head, palpi, antenne, thorax, and abdomen above wainscot-
colour; legs and underside pale and shining; patagia extending
a little beyond thorax. Fore wing greenish buff; three yellow
patches on costa passing into orange and bordered with brown,
extending a little below middle of wing, the outer patch is made
up of two elliptical yellow dots. Hind wing smoky, tinged with
golden and spotted with darker underneath ; fringes paler.
Exp. 31 mm.
Hab, Kebea, Dinawa.
122. NAcoLEIA CHARESALIS W1k.
123. GoNIORHYNCHUS GRATALIS Led.
124. DEBA SURRECTALIS WI1k.
125. D. ALTHEALIS WI1k.
126. BotyoDrEs ASTALIS Guen.
127. B. FuAviBasaLis Moore.
128. SYLEPTA MULTILINEALIS Guen.
129. S. cHRoMALIS WIk.
130. S. cHALYBIFASCIA Hmpsn.
131. S. pissrprrauis Led.
132. S. AURANTIACALIS Fisch.
133. 8. sonitucis Hmpsn.
134. S. PERNITESCENS Swinh.
Proc. Zoot. Soc.—1907, No. VI. 6
§2
135.
136.
137.
138.
139.
140.
141.
142.
143.
144.
145.
146.
147.
148.
149.
150.
151.
152.
153.
154.
155.
156.
157.
158.
159.
160.
161.
162.
163.
164.
165.
166.
ove
168.
169.
MR. G. H. KENRICK ON PYRALID&
SYLEPTA OCHRIFUSALIS Hmpsn.
S. ACRIDENTALIS Hmpsn.
AGATHODES OSTENSALIS Geyer.
GLYPHODES PFEIFFER Led.
. LATICOSTALIS Guen.
. HILARALIS W1k.
. suRALIS Led.
. AMPHITRITALIS Guen.
. UNIONALIS Hiibn.
. CELSALIS WI1k.
. STOLALIS Guen.
. IrysaLis WIk.
. ZELIMALIS W1k.
. EURYTUSALIS WIk.
. cmSALIS WI1k.
CANTHUSALIS WIk.
. BICOLOR Swains.
. ACTORIONALIS WI1k.
. INDICA Saund.
. FLAVIZONALIS Hmpsn.
. UMBRIA Hmpsn.
. AGATHALIS WI1k.
. MARGARITARIA Cram.
AAA DAD a AO a Oam @ oD Pom @
. SUBACUSALIS W1lk.
Q
. SECTINOTALIS Hmpsn
. DELICIOSA Butl.
. OPHICERALIS W1k.
. GLAUCULALIS Guen.
. EXAULA Meyr.
. TRICOLORALIS Pag.
. EZGEALIS Swinh.
. MARINATA Fabr.
. DOLESCHALI Led.
. HYPOMELAS Hmpsn.
QPQeQeaannmm
. EURYGANIA Druce.
[Jan. 15,
MOGI FROM BRITISH NEW GUINEA. 83
170. G. QUADRISTIGMALIS, sp. n. (Plate IV.)
Male with antenne slightly pectinated ; female quite simple.
Head, legs, antenne, and palpi bright ochreous; patagia in
male twice the length of the thorax; thorax and body ochreous ;
tuft black. Fore wing: ground-colour ochreous with slight
purple irridescence ; three equidistant dark purple bands on the
costa are produced until they meet an irregular band of the same
colour from inner margin along the middle of the wing; a faintly
darker subterminal line. Hind wing divided sharply into three
triangular divisions; those on the costa and inner margin
ochreous, but the middle one is iridescent and has a darker
transverse bar; on the hind margin are three double black dots
followed by three single dots all surrounded by metallic scales.
Exp. 32 mm.
Hab. Ekeikei.
171. G. pupicatis, sp. n. (Plate IV.)
Head, thorax, antenne, and abdomen pale ochreous above,
shining white below ; palpi with third joint brown; legs white ;
sides of tuft black. Fore wing hyaline with golden and purple
reflections ; a postmedian angulated transverse line and a faint
mark at end of cell. Hind wing similar to fore wing. Fringes
pale.
Exp. 30 mm.
Hab. Kebea.
172. G. PAUCILINEALIS, sp. n. (Plate IV.)
Head, thorax, and abdomen pale with darker longitudinal lines ;
underside, legs, and palpi paler; antenne brown. Fore wing
crossed by narrow dark bands not parallel and each having a
central yellowish line; the third and fourth touch at costa,
beyond them is an irregular subterminal line broader at the
apex; the spaces between the bands are shining white. Hind
wing with only two bands, forming a triangle with apex at angle
of wing; a darker mark on hind margin.
Exp. 24 mm.
Hab. Dinawa.
173. G. MULTILINEALIS, sp. n. (Plate IV.)
Very much like the above, but the bands are so much wider
that the space left for the white ground-colour is so much reduced
as to suggest a dark wing crossed by numerous white lines which
are not parallel. The details can be better shown in a figure
than described.
Exp. 31 mm.
Hab. Dinawa, Ekeikei.
174. G. PERSPICUALIS, sp.n. (Plate IV.)
Head, top of thorax, and abdomen chestnut; patagia, sides of
abdomen, and underside pure white; tuft black. Fore wing
*
84 MR, G. H, KENRICK ON PYRALIDH [Jan. 15,
golden chestnut, with hyaline white blotches surrounded by
darker lines; a basal triangular spot ; an antemedian oval spot;
a postmedian ovate spot, three small dots between these two; an
apical spot with a line of dots below reaching to inner margin ;
two half spots along inner margin; fringes brown, white at
angle. Hind wing hyaline with marginal dark band intersected
by row of chestnut spots; fringes brown, spotted with white,
Exp. 32 mm,
Hab. Dinawa.
175. G. EXQUISITALIS, sp. n. (Plate IV.)
Head, palpi, and thorax brown ; abdomen in the male grey, in
the female yellow; tuft black. Fore wing chestnut faintly shot
with blue and purple; the spots hyaline and yellowish in the male
but white in the female ; a streak from costa at base sloping to
middle of wing in hind margin; an antemedian triangular costal
spot with long tapering end; a broad postmedian costal spot
extending almost to hind margin, with a darker outside margin
and slight projection into the spot from outside; two very faint
purple, parallel, subterminal lines; fringes golden. Hind wing
pale straw-colour ; in the middle of the hind margin are a few
black dots set in bright golden scales; fringes golden. In the
male the hind wing is dark grey and the fore wing darker.
Exp. 32 mm.
Hab. Kebea.
176. G. BRUNNEOMARGINALIS, sp. n. (Plate IV.)
Head, thorax, and palpi green; antenne dark; abdomen green,
but silvery beneath. . Fore wing apple-green with yellowish costa
and margins; one black dot at end of cell and one in the middle.
Hind wing green, one large black dot at end of cell, margins
having a scorched appearance fading into white nearer the body.
Fringes smoky, but inclined to chestnut at angle.
Exp. 32 mm.
Hab. Kebea.
177. G. LACERITALIS, sp. n. (Plate IV.)
Head, thorax, and palpi apple-green ; antennze and last joint of
palpi brown; legs white, but tibie of fore legs with brown hairs ;
abdomen green above, white at sides and bluish white beneath ;
tuft black with white centre. Fore wing apple-green; hind
margin distinctly and irregularly scalloped, edged with brown as
if scorched ; one black dot in middle and one at end of cell.
Hind wing apple-green, scalloped and scorched like fore wing;
one black dot at end of cell; inner margin white and hyaline
with fringe of same; other fringes brown.
Exp. 44 mm.
Hab. Ekeikei, Kebea, Dinawa, Mafalu.
1907.] FROM BRITISH NEW GUINEA. 85
178. Pygospiua Bivirratis WIk.
179, P. Mareinauis, sp.n. (Plate IV.)
Head purplish grey; palpi, thorax, and abdomen dark grey
above, shining white below; legs white; patagia grey shot with
purple. Fore wing grey shot with purple; a white patch on vein
below costa at two-thirds length of wing from base, 2 mm. wide,
ending in three white dots towards inner margin; a second white
patch between base and first patch; between this and angle of
wing two smaller patches and one still smaller on inner margin ;
fringes grey. Hind wing dark grey shot with purple, discal
portion semitransparent with central spot and serrated dark band;
fringes white.
Exp. 45 mm.
Hab. Mafalu.
180. P. IMPERIALIS, sp. n. (Plate IV.)
Head, thorax, and abdomen grey, striped with white; palpi
black, first and second joints white; underside white. Fore
wing dark grey shot with rich purple; a round white spot near
base surrounded by four white dots; two median, two postmedian,
and two smaller anteterminal white blotches. Hind wing:
ground-colour as fore wing, one long triangular basal blotch, five
other blotches on disc. Fringes dark, flecked with white at angle
of wing.
Exp. 36 mm.
Hab, Ekeikei, Kebea,
181. H&orria viressormpEs Moore.
182. H. pominauis Led.
183. PotyrHuiprA MACRALIS Moore.
184. P. pivaricata Moore.
185. LEPYRODES PERSPICATA Fabr.
186. Merra@a NEBULALIS Wlk.
187. Leucrnopes apicaLts Hmpsn.
188. SAMEODES TRISTALIS, sp. n. (Plate IV.)
Head, legs, antennee, and palpi very dark grey; thorax slightly
paler; abdomen with first three segments paler. Fore wing dark
grey with a V-shaped white transverse mark having its apex on
the inner margin ; near the outer margin of this mark is a thin
angulated dark line. Hind wing dark with irregular transverse
paler band ; fringes dark, spotted with white.
Exp. 24 mm.
Hab. Dinawa.
189. Merrocrena stainronr Led.
190. THLIPTOCERAS OCTOGUTTALE Feld.
&6. MR. G. H. KENRICK ON PYRALID# [Jan. 15,
191. ARCHERNIS CALLIXANTHA Meyr.
192, A. EUcosmA Turner.
193. A. I1GNEALIS WI1k.
194, IsocENTRIS MINIOSALIS Guen.
195, I. mrntauis Warr.
196. Maruca TESTULALIS Geyr.
197, PacHyNoa THOOSALIS W1]k.
198. P. sprLosomorpEs Moore.
199. PACHYZANCLA DESMIOIDES Hmpsn.
200. P. PALLIDALIS Hmpsn.
201. PRogDEMA INSCISALIS WI1k.
202. BaoraRCHA HYALINALIS Hmpsn.
203. B. timpata Butl.
204, CALAMACHROUS TRANQUILALIS Led.
205. C. ALBIPUNCTALIS, sp. n. (Plate IV.)
Palpi dark purplish brown ; head orange; thorax and patagia
purplish brown ; abdomen above fuscous; thorax and abdomen
below, legs and underside of palpi white. Fore wing dark purplish
brown, with a subtriangular patch from the middle of the costa to
middle of the wing orange, paler inside and with a minute orange
dot in the middle; hind margin broadly orange with a row of
subterminal dots. Hind wing white on costa, the middle of the
wing occupied by a triangular purplish patch, the apex reaching
the these of the wing ; between this and the inner margin 1s walnihe
the hind margin broadly orange and fringe orange; in the centre
of the patch are two minute white dots edged with dark.
Exp. 44 mm.
Hab. Dinawa.
206. PIONEA ABLACTALIS Wlk.
207, P. nypstusALis WIk.
908. P. ALBIFIMBRIALIS Wlk.
209. P. BRACTEALIS, sp.n. (Plate IV.)
Head, thorax, abdomen, and legs pale ochreous, inclining to
reddish above but shining white below, very noticeable on tibia ;
palpi with third joint chestnut. Fore wing golden yellow; costa
margined with reddish; five equidistant, thin, angulated, some-
what interrupted transverse lines, Hind wing the same, with
three faint lines. Fringes pale.
Exp. 30 mm.
Hab, Dinawa, Ekeikei.
1907. | FROM BRITISH NEW GUINEA. 87
210. Pyrausta CELATALIS WIk.
211. P. ERtoprsatis W1k.
212. P. ceapEsaLis WIk.
213. P. rriricatis, sp. n. (Plate IIT.)
Head and thorax orange; patagia darker; abdomen paler with
chestnut markings along the crest; palpi and antenne dark
brown; underside and legs whitish; tarsi ringed with darker.
Fore wing orange with dark margin on the costa, dark nervures,
and the cell margined with darker ; an oblique line, dark brown,
passes through the cell and reaches the hind margin; a second
oblique dark line reaches from the middle of the hind margin to
near the apex. Hind wing straw-colour, margined with darker.
Fringes smoky with white edges.
Exp. 26 mm.
Hab. Dinawa.
214. PoLYGRAMMODES GROSSALIS Guen.
215, P. puRPURALIS WI]k.
216. OmpHisa INGENS Hmpsn.
In concluding this list I desire to tender my most hearty thanks
to all those who have assisted in determining the species and
especially to Sir G. Hampson, Col. Swinhoe, and Dr. Dixey, who
kindly allowed me to see the Walker types in the Hope Museum
at Oxford.
EXPLANATION OF THE PLATES.
PratE Il. Prare IV.
No. No.
6. Phycita dinawa, p. 69. 5. Monodonta passalis, p. 69.
7. Htiella fuscalis, p. 69. 58. Parthenodes rectangulalis, p. 77.
14. Macalla unipunctalis, p. 70. 68. Dracenura unicoloralis, p. 77.
15. na caradriniformis, p. 70. 69. Piletocera inconspicualis, p. 78.
16. “ pomalis, p. 70. 80. Agrotera semipictalis, p. 78.
17. = curtulalis, p. 70. 86. Rhimphalea lineatis, p. 79.
18. i apicalis, p. 71. 97. Ulopeza cruciferalis, p. 79.
20. 33 perdentalis, p. 71. 105. Caprinia castanealis, p. 80.
21. - porphyrealis, p. 71. 106. 45 ocellalis, p. 80.
23. Polylophota truncalis, p. 72. 120. Dichocrocis pardalis, p. 81.
24. Locastra castanealis, p. 72. 121. > bimaculalis, p. 81.
29. Stericta flammealis, p. 72. 170. Glyphodes quadristigmalis, p. 83.
30. 3 cornucalis, p. 73. iil . pudicalis, p. 83.
31. cs subviridatis, p. 73. 172. is paucilinealis, p. 83.
33. Orthaga polychroalis, p. 73. 173. oo multilinealis, p. 83.
34, es chionalis, p. 73. 174. = perspicualis, p. 83.
35. S fuscofascialis, p. 74. 175. s exquisitalis, p. 84.
36. D columbalis, p. 74. 176. i brunneomarginalis,
42. Tipuliforma triangulalis, p. 75. p. 84.
45. Vitessa griseata, p. 75. 177. 53 laceritalis, p. 84.
55. Aulacodes lunalis, p. 76. 179. Pygospila marginalis, p. 85.
56. D purpurealis, p. 76. 180. 35 imperialis, p. 85.
57. % bipunctalis, p. 76. 188. Sameodes tristalis, p. 85.
67. Dracenura torridalis, p. 77. 205. Calamachrous albipunctalis, p. 86.
108. Tyspanodes radiata, p. 80. 209. Pionea bractealis, p. 86.
213. Pyrausta triticalis, p. 87.
88 DR. E. A. GOELDI ON MARMOSET [Jan. 15,
5. On some new and insufficiently known Species of Marmoset
Monkeys from the Amazonian Region. By Prof. {Dyes
Bur A. Gorupt, C.M.Z.S., Director of the Para Museum.
[Received November 23, 1906. |
(Text-figures 20-23.)
On the occasion of the Sixth International Zoological Congress,
held at Berne (Switzerland) in August 1904, I presented a paper,
‘Nova zoologica from the Amazonian Region, dealing especially
with new Vertebrates,” in which I discussed at some length new
and little-known representatives of the family of Callitrichidee
(Hapalide, auctorum*) from the Upper Amazon, especially from
the Rio Purtis, as follows:—(1) A species of J/idas whose close
relationship with Midas rufiventer Gray I recognised at first
sight, and to which later on, after comparing it with the
type-specimen of this latter in the British Museum in London
(at the time of my visit to the International Ornithological
Congress in July 1905), I decided to give the name of Midas
griseovertex. (2) A second species of J/idus, evidently related to
M. labiatus Geoftr. (1812), MW. illigert (id. 1845), and M. weddelli
(id. 1849), and having as its most characteristic distinguishing
feature an enormously long white moustache, which afterwards,
on the same occasion, I decided to name J/idas imperator.
(3) Midas pileatus, described by Geoffroy in 1848 from the original
specimen from the Rio Javary, kept in the Paris Museum, and
until recently not represented in any other Museum, so far as
could be judged from current zoological literature. I was then
able to show a splendid pair of this very rare species. (4) Midas
fuscicollis, described so long ago as 1823 by Spix, but only from
immature specimens, the habitat of the adult animal of both sexes
having escaped notice, as it appears, in a surprising manner until
1904. (5) Midas mystax Spix, originally described from the Rio
Solimoés, represented in my exhibited collection by a very dark
specimen from the Rio Jurua.
One of the essential conditions for arriving at certainty in my
conclusions, as above suggested, was the careful examination of
certain individuals kept as type-specimens in the British Museum.
The purpose of the present article is principally to record the
result of my investigation in this respect, which proves to be most
interesting and instructive. To say it at once: the most remark-
able and unexpected discovery was the fact, that of the two
specimens of Midas rufiventer, type and quasi co-type, on the
shelves in the British Museum, the latter does not coincide with
the former, but, on the contrary, should constitute the type-
specimen of a new, overlooked, and undescribed species of the
genus JJidas.
* See Thomas, Ann. Mag. N. H. (7) vol. xii. (1903).
1907. } MONKEYS FROM THE AMAZONIAN REGION. 89
Minas THOMAST, sp. n. (1905).
[Labelled in the British Museum: “ Midas rufiventer, 3,
‘The red-bellied Tamarin, Upper Amazonia. W. Bates.
Exp. 1857, Tunantins, north side of Amazons.” |
Dorsau Aspect.—Anterior half of a dark, deep blackish-brown
colour (somewhat like the half-grown J/, fuscicollis), extending
from behind the ears as far as the middle of the back. Hinder
half of the same brown, the tips of the hairs being however
mottled with a lighter greyish-brown, but in a much less pro-
nounced degree than in the typical JL. rufiventer.
Head (text-fig. 20) black, with an insignificant median small
light spot beginning just between the ears, behind the vertex ;
a still less noticeable lightish dash, consisting only of a few
greyish hairs, almost forming an anterior continuation, is per-
ceptible in the same median line just in front of the former.
Details of the region of mouth and nose as in the type of
M. rufiventer.
Text-fig. 20.
Back view of head of Midas thomasi.
Arms dark blackish-brown, passing into a true black towards
the hands, not being, however, a brilliant black as in the
M. rufiventer type.
Legs. Yixterior side of thighs of the same colour as the posterior
half of the back, turning to black toward the feet.
Tail black.
VENTRAL ASPECT.—Anteriorly a cross-zone of light reddish-
yellow (viz., inside of arms and a band across the breast as wide
as the insertions of thearms). Rest from there backwards reddish
rust-yellow (lighter than in the new species MV. griseovertex G.,
and much lighter than in J/. rufiventer type).
The light under side is abruptly separated in the region of the
neck and breast from the black of the neck and head. (This
light abdominal colour cannot be attributed to fading of an old
90 DR. E, A. GOELDI ON MARMOSET [Jan. 15,
Museum specimen, perhaps unduly exposed to light on the
shelves, because in that case such change would certainly be even
more noticeable in the colouring of the back, which, although
more exposed, still holds however its colour fast ; consequently
the impression is produced of a decidedly distinguishing character
of specific rank.) The light abdominal colour extends to the
under side of the insertion of the tail, as in J. rufiventer type
and in the other new species, JZ. griseovertea.
On separating the fur the deeper portion is of the dark
colouring, which is lighter in the anterior than in the posterior
half of the body, and thus corresponding perfectly with the scale
of colouring of the external aspect of the fur.
The only existing specimen (stuffed) dates, as noted above in
the copy of the label, from Tunantins, Upper Amazon, where it
had been collected in 1857 by Henry W. Bates.
Let us now proceed to the description of the type-specimen
of the true Midas rufiventer Gray, m the British Museum, in
order to relieve once for all the deplorably embarrassing situation
caused by the very defective descriptions given up to the present
time.
MIDAS RUFIVENTER.
[‘‘ Brit. Mus. Reg. 43.10.12.6. Type (skin). Locality :
South America. Purchased of Argent. ¢ (skull).”]
I must state at the outset, that I was informed by Mr. Oldfield
Thomas that the specimen had been obtained from a dealer, a
fact which accounts for the vagueness of the locality.
In general the results of my examination coincide with the
description given by. Gray in his ‘Catalogue of Monkeys, ete.’
1870, p. 66, of this original specimen, only that it is too brief.
DorsaL ASPECT.—Predominating colour of the back a dark
blackish-brown. Hairs light greyish for the apical + of their
length. Anterior part of back less greyish, posterior part. of
back more so. The fur down to the roots dark brown (not
light greyish-brown).
Head (text-fig. 21). General colouring of the head a deep
black, not blackish-brown. Mouth bordered with sparse whitish
hairs. Central zone between and beneath the nostrils sparsely
covered with lightish grey hairs (not brilliantly white).
A spear-shaped dark rust-red median stripe extending from the
vertex between the ears downward to the insertion of the nose,
the rust-red beg of the same intensity as the colour of the whole
abdominal side. From the vertex backward, in form of a double
ploughshare, over the occiput down to the neck spreads out
a patch of a greyish-brown colour, which combined with the lance-
shaped frontal marking forms a comparatively broad triangle with
a tendency to lateral posterior development.
Arms. Outer side deep dark brown (like the head), slightly
lighter on the upper arm.
1907. ] MONKEYS FROM THE AMAZONIAN REGION. 91
Legs. Outer side of the same colour as the posterior part of
the back.
VENTRAL ASPECT.—General colour a deep dark chestnut reddish-
brown, considerably darker than in the Purus species, J/. griseo-
vertex.
Tail very long, dark blackish-brown, turning blacker toward
the end.
Text-fig. 21.
Back view of head of IWidas rufiventer.
So far as I know the only figure as yet published of a specimen
of Marmoset Monkey, closely. resembling this ty pe-specimen of
Midas rufiventer in the British Museum, is that in the supple-
mentary plate 36 of the work of Reichenbach, under the name
of “ Midas erythrogaster Natterer, Mus. Vindob.,” made, as stated
on p. 14, from an original sketch by a Mr. T. E. Zimmermann
of a skin collected by Natterer and preserved in the Vienna
Museum. With no little surprise I discover, however, that
Pelzeln, the excellent monographer of the Brazilian mammals
collected by Natterer, enumerates this very specimen, and also
the figure in Reichenbach, as synonyms under the head of J/das
labiatus Geoffroy, a procedure which seems to me more than
doubtful, not to say flatly erroneous. In this figure, though
worthy of criticism in some respects (the most salient defect
being the substitution of the delicate mottling by rough. daubs of
‘ieee on a greyish ground), one essential feature nevertheless
stands out clearly, namely the rust-coloured frontal spot, coinciding
perfectly with the abdominal colour.
I have never been able to consult either the full description,
or any figure, or any other useful information concerning the
M, elegantulus Slack, Proc. Academy of Natural Sciences of
Philadelphia, 1861, p. 463, always quoted as a synonym of
M. rufiventer Gray.
92 DR. E. A. GOELDI ON MARMOSET [Jan. 15
Having now finished these explorations of the treasures stored
up in the British Museum as an essential preliminary and
solid basis for the discussion of the respective questions that arise,
I will proceed to the detailed description of the two new species
of Marmoset Monkeys of the Purts Region, mentioned at the
beginning of the present article. -
MIDAS GRISEOVERTEX, Sp. 0.
In general aspect closely similar to I. rufiventer Gray and
M. thomasi Goeldi, in the sharp contrast between the bright
rusty-coloured abdominal side and the dark colouring of the
dorsal aspect, but distinguishable at a glance by the greyish-
white, good-sized rounded frontal patch.
Among the collections made in the Purts and Acre Regions
by our Museum Expeditions (1903-1904) there are seven indi-
viduals of this most interesting and well characterised species of
Marmoset Monkey (skins, skulls of all and trunks of some),
4 being malesand 3 females. One mounted pair (¢ @ ) remained
in the Museum at Berne, labelled as above; three mounted indi-
viduals (¢, 2, and a half-grown young one) are kept in the Para
Museum ; and the remaining pair of skins (¢ @ ) are intended for
the British Museum.
Dorsau Aspect.—General colour deep blackish-brown, excepting
head, hands, feet, and tail, which are positively black. This
colour remains pure from the nape of the neck backward for
one-third of the length of the back. The hinder two-thirds of
the back shows a mottling due to the light greyish tips of the
hairs, which terminal points measure about + of the entire length
of the hair and are slightly longer toward the hips; the intensity
of the mottling increasing gradually backwards, being most pro-
nounced in the sacral region, presenting even a whitish appearance,
when seen from certain oblique directions.
On separating the fur in the region of the shoulders, the
impression of colour is in general the same as the exterior, that
is deep blackish-brown, excepting that the lower third of the
hairs towards the roots forms a slightly lighter zone, especially
laterally to the median line. Making the same examination in
the region of the hips, the light zone is scarcely apparent, the fur
being of the general colour almost down to the roots.
Head. General colour sooty-black. A narrow zone of whitish
hairs bordering the whole extent of the mouth. In the region
of the upper lip the white zone rises in a very conspicuous
triangular zone, with its broad base resting on the circular white
band of the mouth. One very noteworthy feature of this
triangular zone is that the lateral oblique lines cross exactly
the middle of the nostrils, so that the exterior half of each
falls in the blackish region, the interior half in the whitish
triangle.
At the vertex, from a line connecting the anterior borders
of the ears, commences a whitish patch (text-fig. 22), occupying
1907.] MONKEYS FROM THE AMAZONIAN REGION. 93
about 3 to 2 of the width between the ears and widening slightly
backwards, but in no specimen acquiring the double ploughshare-
shape of the rust-coloured patch of the Midas rufiventer type. In
young individuals this patch is rather more greyish, while in
more aged specimens it becomes nearly white. In length it
extends as far as the posterior border of the occipital region.
Arms. Outer side from shoulder to elbow of the same sooty
blackish-brown as the whole shoulder region; fore-arms and
hands decidedly black.
Legs. Outer side of the same intensely mottled colour as the
entire hip-region as far as to the foot; the foot itself of the same
black as the hand.
Back view of head of Widas griseovertex.
VENTRAL ASPECT.— General colour light reddish rusty, decidedly
lighter than in Midas rufiventer. It embraces the whole area
from the middle of the. throat, breast, and belly, inner side of arms
and legs, and a short distance beyond the insertion of the tail on
its under side (some 5 cm.). On the flanks the dorsal and ventral
colours are very abruptly separated, it being noteworthy that the
rusty-coloured hairs of the abdominal side are only about half as
long as the adjacent dark-coloured hairs of the dorsal part.
Tail long, exceeding the length of the body by that of the head.
Colour the same black as of the fore-arms and hands, but here
and there with rows of hairs with light brownish tips, which may
be interpreted probably as the vestiges of an annular arrangement.
MIDAS IMPERATOR, Sp. 0.
In general aspect, especially in colour, somewhat similar to
M. labiatus Geoftr., MW. uligert, and M. weddellii, but easily dis-
tinguished by the immense white moustache, which becomes
94 DR. E. A. GOELDI ON MARMOSET [Jan. 15,
phenomenal especially in the adult male. (Text-fig. 23 represents
the head of an old male individual.)
Ow: Purts collections embrace five skins (and skulls) (395 ¢,
2 2 2), two from the Rio Acre and three from the upper Rio
Puriis. <A family of three individuals (¢ 2 adults, and a young
one) are already mounted in the Par&é Museum; the two
remaining (¢, @) skins, of half-grown specimens, are destined
to be sent to the Museums of Berne and London respectively.
This species is distinguished by a more or less pronounced
reddish greyish-brown general colour of the dorsal side, light rust-
coloured abdominal side, dark feet, hands, and head, excepting
the white cireumbuccal zone and the skull-cap, which tend to
become grey with age.
Head of Midas imperator.
Dorsau aspect.—The reddish-rust colour is more conspicuous in
the above mentioned old male, and is already noticeable in the
two young males (one stuffed in the Para Museum and one skin).
General colour as above stated ; it may be noted, however, that the
greyish tinge, in the same way, is more pronounced in the female.
On separating the fur nearly every one of the five specimens
presents a slightly different appearance. In the two adult stuffed
specimens in the Para Museum the lower half is uniformly dark ;
the outer half shows alternately a light zone, then a dark one,
followed by a light one and a dark one, the terminal zone being
again lighter, making five zones of equal breadth in the outer half
and giving the general impression of two equal pairs of light and
dark zones.
1907. | MONKEYS FROM THE AMAZONIAN REGION. 95
Examining in the same manner the two skins of half-grown
individuals (2, ¢), we see immediately, that while the lower
half is dark as above, the outer half shows alternately a light
zone, then a dark one, and finally the terminal lighter one, giving
as a total result the impression of only one complete pair of
light and dark zones. The female skin presents nevertheless a
peculiarity in the circumstance that near the roots of the hairs
there is an unmistakable trace of a basal light zone—a peculiarity
of this one individual among all the five known.
Proceeding now to the examination of the same details in
the quite young stuffed specimen of the mounted family in the
Para Museum, the general impression is of only one well-marked
pair of dark and light zones, from within outwards, while on closer
inspection the second outer paix noticed in the half-grown speci-
mens is unquestionably recognisable.
Recapitulating these facts, the general conclusion is that the
number of light and dark zones tends to increase with age. We
have here one more useful hint, that in the discrimination of species
prudence is required in judging the value to be attributed to
such variations in colour, which in the present case can be fairly
proved to be due to differences of age and sex.
Head. All parts of the full front view of the face and direct
lateral view are sooty-black in the adult female, the quite young
individual (g) and the two skins (¢ 2), while the old male
shows the face somewhat mixed up with greyish-brown hairs.
The circumbuccal zone, however, is again an exception from
this general colour, being saliently pure white. This zone is
relatively broader than in any of the preceding species, including
the entire nostrils in the old male, while in the other four speci-
mens the border-line of the white colour curves below the nostrils,
arising again on the ridge of the nose at about one-third of its length.
The same white colour characterises the phenomenal moustache,
which already in infant specimens is of a greater length than in
any other of the moustached species known to me. The old
male (text-fig. 23), however, beats the world’s record, the longest
hairs attaining a length of no less than 57 to 60 mm.!
My collectors inform me that the moustache is worn in life not
straight, but with the modern upward twist. The two oldest
specimens show the strange strong development of the white hairs
of the circumbuccal zone extending even over the whole area of
the lower jaw, not including the chin and inferior side of the jaw.
As a result, the old males appear bearded as well as moustached.
In the three younger individuals the crown of the head is
brownish-black, with a tendency towards forming the already
familiar favourite lighter-coloured patch of a pale brownish shade.
In the adult female the patch is diamond-shaped, almost dirty
greyish-white, and therefore very conspicuous. In the old male
a pronounced greyish cast spreads all over the top of the head,
without forming any distinct patch.
Arms. Outer side of upper arm the same colour as the back,
96 DR. E, A. GOELDI ON MARMOSEL | Jan. 15,
but a little lighter in all specimens. Fore-arms and hands
gradually darker towards the extremity. This is especially the
ease in the three young specimens and the old female, while the
male shows on the outer side of the fore-arm and hands the same
dark greyish cast as noted above in the description of his face.
Legs. Outer side of thigh the same colour as the back, but as a
rule a little lighter, especially m the old female. The outer
side of the foot is of the same dark colour as the hand.
VENTRAL ASPECT.—The colour forms a marked contrast with
that of the dorsal aspect, by having a pronounced tendency towards
a rusty red, especially in the younger specimens. This rusty-red
shade covers the whole under side, from the throat backward in-
cluding the inner side of the arms and legs and no small extent of
under side of tail. The same colour tends to form regions or
patches of a deeper shade, one between the arms, covering the chest,
and another between the legs, covering the abdomen and sur-
rounding the anus. The chest-patch, especially in the old female,
looks as if soiled by dried blood.
Tail. The tail is long, nearly half as long again as the body
(the average length of four specimens measured being as 11 : 8).
In the younger specimens the dorsal side nearly from the inser-
tion to the end is of the same dark colour as the feet, while the
lower side shows a gradually narrowing line of the above mentioned
deep rusty red fading toward the extremity. In the adult
specimens the colour of the tail tends to become uniform in its
whole circumference. In the old female the upper side is darker
than the lower at least for two-thirds of its length, the terminal
third being of a dark grey, due to the black roots and the ght tips
of the hairs. The under side for about half its length has the general
hight rusty-red colour of the ventral aspect, the terminal half
being alike on the dorsal and ventral side, a dark grey.
Most aberrant is the colouring of the tail in the old male.
From the very insertion the bright rusty-red colour predominates
in its whole circumference throughout its entire length with the
exception of a dark terminal tuft. The rusty red is most
noticeable to a certain extent (one-eighth) from the insertion
backward. Approaching the end the colour is more mixed with
dark, owing to the greater extent of black at the roots of the
individual hairs. <A’ darker isolated patch exists also on the
second eighth.
MIDAS FUSCICOLLIS Spix.
This species, it is true, was long ago described, having been
introduced into science in 1823 by the Bavarian explorer J. von
Spix from individuals obtamed on the Rio Javary, but it is
evident from our much richer material (eight specimens, 6 ¢ ¢
and 2 2 2), collected on the Rio Purtis, that the pelage of
the full-grown animal has never been properly described and
figured. Referring to a comparison of the two figures 3 (adult
coat) and 3.@ (juvenile coat) of my original coloured Marmoset-
HOOT: MONKEYS FROM THE AMAZONIAN REGION. 97
Monkey plate, presented to the Zoological Congress at Berne and
now being reproduced for a separate publication, it is especially
noticeable that the adult coat is distinguished by the deep blackish-
brown colour of the anterior part of the body.
Briefly described, the colour of a typical adult male is as
follows :—
Dorsal aspect of anterior part of body, as far back as behind
the shoulders, of a uniform sooty brownish-black. On separating
the fur, two-thirds of the length of the hairs is lighter, greyish
or dirty white near the roots, pale greyish-brown outwards, the
outer third shading into the above-mentioned brownish-black,
without presenting any tendency to zonal arrangement. ‘The
posterior part of the body presents a mottled appearance, due to
the intervention of yellowish-reddish zones. On separating the fur
in this region, two-thirds of the length of the hair is quite uni-
form dark; the last third is about equally divided between the
light zone and a black terminal one.
Singularly deep rusty-red are the rump and thighs. This is
due to the absence of the dark terminal zone of the hairs in
this region, each hair being black at its base, and the terminal
third entirely rusty. Tail, arms, and feet black.
Dorsal and ventral aspects are noticeably divided by a lateral
rusty-red stripe of shorter hairs on each side, running from the
arm-pit to the flank. The real median ventral stripe, wider than
the just-mentioned lateral one, again assumes the dark brown
colouring of the anterior part of the body.
In the face the most salient features are the white eyebrows,
meeting in the median line at the base of the nose, forming
a very striking double crescent. Circumbuccal zone whitish,
more extensive than in all the preceding species, but not so
sharply outlined.
Minas PrLEAtus Is. Geoffroy et Deville.
This species was figured in 1848 by Geoffroy in the ‘ Archives du
Muséum,’ vol. v. pl. 31, and described under the name of “ Tamarin
a calotte rousse,” p. 569. Up to the time of my paper read at
Berne, at the International Zoological Congress, it seemed to me,
judging from the literature within my reach, that it was repre-
sented only by the single original specimen, coming from the Rio
Javary, and preserved in the Paris Museum. (I saw it there
some days before, without a label, stowed away on a side-shelf.)
At that time we had obtained from the upper River Purtis two
other specimens, a pair. This splendid species has its chief
distinguishing marks in the cinnamon-coloured sealp-patch, the
brownish-black general colour, and the sharply outlined white
circumbuccal zone, including the nostrils, which are completely
surrounded by a narrow white band.
Being in London for the Meeting of the Fifth International
Ornithological Congress in July 1905, Mr. Oldfield Thomas, of the
Proc. Zoou. Soc.—1907, No. VII. 7
98 DR. E. A. GOELDI ON MARMOSET [Jan. 15,
department of mammals at the British Museum, to my no small
surprise showed me a small series of skins from different points
in the Upper Amazon region, one having been furnished by
Dr. von Ihering, from the River Jurua expedition.
After my return to Para I saw by the ‘ Revista do Museu
Paulista,’ vol. vi. (1904), p. 416, which had been sent in my
absence, that Dr. von Ihering refers to three individuals in the
collections made by Mr. Garbe at the time of the above men-
tioned Jurud trip. He, however, classifies it as a new subspecies
—WM. pileatus jurwanus—which according to his statement is
distinguished by the dark colour of the hair of the back, which in
the description of the original type-specimen, made by Geoffroy,
is stated to be reddish at its base. It is true, that Geoffroy expresses
himself in the following terms :—‘“ Le corps est supérieurement re-
couvert, ainsi qu’on l’observe si souvent chez les Hapaliens, de
poils roux dans la plus grande partie de leur étendue, anneleés de
blanchdtre et de noire vers la pointe. Il] resulte de cette dis-
position, chez JZ. pileatus, un mélange de gris et de noir, mais
non des bandes alternatives de l’une et de autre couleur.”
Although the adjective “roux” may not be perhaps a happy
term, the rest of the description and the figure itself indicate
an animal which does not differ essentially from the specimens
examined by me in London, nor from my two mounted speci-
mens here at Para. Geoftroy evidently wrote with the desire to
differentiate the uniform general colouring of this species (which,
as already seen, is usually a common feature of most Amazonian
Callitrichidee) from the distinctly striped colouring of certain
southern Marmoset Monkeys (Hapale jacchus, &c.).
On separating the fur of the back of my two individuals in the
Para Museum, the hair appears to be of the same deep blackish-
brown as in the case of WW. griseovertex. The terminal fourth
then shows a light greyish-white zone, followed by a terminal
nearly black one. It follows therefore that in this detail my
Purts specimens accord with the Jurua specimens of Ihering.
However, this detail does not seem to me of sufficient weight to
justify the establishing of a separate subspecies.
MipAs MyYsTAX Spix.
Acquaintance and description of this species date from the
same period and come from the same source. Spix brought his
specimens from the River Solimoes; a female is figured on plate 22
of his work. Our Museum possesses a considerably darker male
specimen from the River Jurua.
Instead of entering into a detailed description, I would simply
call attention to the characteristic fact, that the white circum-
buccal zone is nearly identical im extent and form with that of
the preceding Midas pileatus, but with a stronger tendency to
form a moustache.
1907. ] MONKEYS FROM THE AMAZONIAN REGION. 99
Mipas 1LLicert Pucheran.
I have recently received, through the kindness of the wife of
the German Consul at Para, Dr. Olshausen, an example of another
species of Amazonian Marmoset Monkey hitherto unrepresented
in our collections. It had been obtained at a very early age from
an Indian woman at Iquitos, and kept as a pet until its mistress
was about to sail for Europe. It lived several months at the
Museum until about half-grown, and is now mounted.
Dorsau Aspect.—Principal colour a lovely dark brown-red
covering the nape of neck, shoulders, and outer side of arms and legs,
embracing with a small band the rump and base of the tail. The
back, however, properly speaking, forms an exception to this colour,
bearing a very long oval patch, distinctly outlined, of dirty black
in the centre and greyish borders, with the tendency to form
posteriorly dimly apparent light and dark transverse bands. Head,
hands, feet, and tail black.
VENTRAL ASPECT.— Uniformly brownish-black from the throat
as far as the anus, embracing the inner side of arms and legs,
separated from the dorsal patch by a brownish-red margin.
In the black face we again find the white circumbuccal zone.
But in this case it leaves the nostrils free and the white runs up
to the cheeks in a triangular form nearly as far as the outer
corner of the eye.
By this feature the animal immediately proves to belong to
the group 6, subdivision a of the classification of Schlegel (‘ Singes,’
p. 262), that isto say tothe Hapale devillei group. Among all the
figures I have at my disposal, my specimen corresponds best with
the animal represented in plate 13 of the ‘ Proceedings of the
Zoological Society,’ 1871, by Bartlett, with the designation of
Midas devillei $ (considerably better than with the figure of
M. devillei, fig. 3, plate 6, in the ‘ Atlas’ of Castelnau, which lacks
any trace of a distinctly coloured dorsal patch). As the animal
there represented is attributed by Schlegel (‘ Singes,’ p. 263) and
by Forbes (‘Handbook of Primates, p. 145 seq.) to Hapale
illigert Pucheran (‘ Revue de Zoologie, 1845, p. 336), and as the
description of this species given by these two authors coincides
satisfactorily with my Iquitos individual, I think I have to do
with a half-grown specimen of Midas illigeri Puch.,a Marmoset
Monkey stated to be fairly abundant in the Peruvian Amazons.
7*
100 PROF. E. RAY LANKESTER ON THE | Feb. 5,
February 5, 1907.
His Grace Tue Duke or BeprorD, K.G., President,
in the Chair.
Mr. F. Martin Duncan, by permission of the Charles Urban
Trading Co., Ltd., gave a cinematograph exhibition of animals in
the Society’s Gardens and other zoological subjects, chiefly the
life-history of Insects.
Mr. Oldfield Thomas, F.R.S., F.Z.S., exhibited a collection of
Mammals and Birds from the Islands of Saghalien and Hokkaido,
N. Japan, made by Mr. Malcolm P. Anderson in carrying out the
Duke of Bedford’s Exploration of Eastern Asia. Mr. Thomas
proposed to give a full account of the Mammals on a later
occasion.
Dr. W. T. Calman, F.Z.S., read a paper entitled ‘On new or
Rare Crustacea of the Order Cumacea from the Collection of the
Copenhagen Museum. Part I. The families Bodotriide, Vaun-
tompsoniide, and Leuconide.” b
This paper will be published entire in the ‘ Transactions.’
The following papers were read :—
1. The Origin of the Lateral Horns of the Giraffe in Foetal
Life on the Area of the Parietal Bones. By H. Ray
lankestprs) WIA. DiSe., use, SEeRES* HeZzas:,
Director of the Natural History Departments of the
British Museum.
[Received February 5, 1907. ]
(Text-figures 24-36.)
A remarkable and wide difference between the Giraffe and the
Okepi is constituted by the position and relation of the lateral
horns in these two animals in regard to the bones of the skull.
AsI pointed out in my memoir on the Okapi read in 1901 (Trans.
Zool. Soc. vol. xvi. p. 279), the bony horn-cone of that animal is
attached to the frontal bone, and it is the frontal bone which is
raised into a boss for its support, whilst even in the hornless skulls
supposed to be those of the female these frontal bosses are present.
On the other hand, in the young Giraffe the main axis of the
lateral “ossicone”* falls within the area of the parietal bone
* T use the term “‘ ossicone ” in the present paper for the independently ossifying
bony cones which are found in Okapi and Giraffe on the frontal and parietal areas
and in the Giraffe also in a median position. In my memoir of 1901 I spoke of such
structures as “ossicusps,” a term which I now wish to apply more generally,
reserving the term “ossicone” for the peculiar separately ossifying cones of the
Givraftide.
1907. ] HORNS OF THE GIRAFFE. 101
(text-fig. 24). The wide-spreading base of the cone-like ossicusp
subsequently encroaches, it is true, over a large portion of the
frontal bone. In the adult both the parietal and the frontal are
Text-fig. 24.
Lateral view of the skull and lower jaw of a very young Giraffe, measuring
30°8 centimetres from the occiput to the anterior border of the premaxilla:
preserved in the British Museum. The drawing is five-twelfths of the natural
size.
oce., occipital crest; gé./., Giraffine conical tumescence of the parietal bone, above
which is developed the lateral ossicone (p.oss.); sfp., the fronto-parietal suture;
ot.l., position of the lateral tumescence of the Okapi, ‘absent here ; gt.m., position
of the median frontal tumescence of the Giraffe, which in this young specimen
is still entirely undeveloped ; o¢.m., position of the median tumescence of the
Okapi’s skull (basinasal) ; pln, prelacrymal vacuity; can., bifoliate canine
(deciduous dentition).
(From Trans. Zool. Soc. vol. xvi. p. 293.)
enlarged and tumescent, and both enter largely into the com-
position of the lateral horn of the adult Giraffe (text-figs. 25 & 33).
The whole form of the skull is rendered different in the two
genera by this relationship of the ossicone to the frontal exclusively
in the one, to the parietal primarily but not exclusively in the
other (see text-figs. 26 & 27).
In a skull of a very young Giraffe (text-fig. 24), probably about
a year old, preserved in the British Museum, the lateral ossicone
is seen to rest almost entirely on the parietal. A transverse
section (text-fig. 28, p. 104) shows that the anterior margin of the
enlarging base of the bone constituting the ossicone has spread—
102 PROF. E. RAY LANKESTER ON THE | Feb. 5,
Text-fig. 25.
Lateral view of the skull of a Giraffe, about two-thirds grown.
oce., occipital crest ; p.oss., parietal ossicone (epiphysis) overlying the parietal conical
upgrowth and spreading on to the frontal bone; sfp., fronto-parietal suture ;
o.t.l., position of the lateral frontal tumescence in the Okapi, absent here ;
g.t.m., the characteristic median tumescence of the Giratte’s frontal, devoid in
this specimen of any secondary cap or epiphysis, absent in the Okapi; pl.v., the
prelacrymal vacuity.
(From Trans. Zool. Soc. vol. xvi. p. 284.)
Text-fig. 26.
(
(yyyt 4
= ihe Le
NY
) vt iM) GH,
in tat
View from above of the fronto-parietal region of the skull of a very young Giraffe.
The parietal epiphyses were already ossified, but separable, and are here removed.
occ.h., exostosis of the occipital crest, which in some adult Giraffes forms a second
pair of “horns” (five-horned Giraffe) ; gtl., the Giraffine lateral tumescence, seen
here to originate in the parietal bone, which carries the conical ossicone aud forms
the Giraffe’s paired “horns ”’; o¢./., the position of the Okapi’s lateral tumes-
cence of the frontal bone, absent in Giraffe; zy., zygomatic arch; po., posterior
angle of the orbit ; ao., anterior angle of the orbit; pl.v., prelacrymal vacuity
sof., supraorbital fossa; sfp., fronto-parietal suture.
(From Trans. Zool. Soe. vol. xvi. p. 291.)
1907. | HORNS OF THE GIRAFFE. 103
invaded as it were—the area of the frontal bone to a very slight
extent.
It seemed hardly possible to doubt that the ossicone of the
Giraffe takes its origin within the area of the parietal bone, but
that conclusion was forbidden by the explicit statement of the late
Sir Richard Owen who, in a paper published sixty-seven years ago
(1840) in the ‘ Transactions’ of the Zoological Society, described a
newly-born Girafte which had died in the Gardens of the Zoological
Society. Owen there states that he found the lateral horns of this
Giraffe to be definitely attached to the frontal bone, and to that
Text-fig. 27.
MATA
Lae
View from above of the fronto-parietal region of the skull of an immature Okapi.
oce.h., angle of the occipital crest ; g.¢.J., position of the lateral tumescence of the
parietal which supports the paired ossicones of the Giraffe, absent here; 0.t.l.,
the Okapian tumescence of the frontal which supports the paired ossicones of
the Okapi; zy., the zygomatic arch; po., posterior angle of the orbit; ao.,
anterior angle of the orbit ; pl.v., prelacrymal vacuity; ofm., the slight median
tumescence of the base of the nasals of the Okapi; sof., supraorbital fossa;
sfp., the fronto-parietal suture.
For comparison with text-ficure 26.
(From Trans. Zool. Soe. vol. xvi. p. 290.)
bone exclusively. He gives the drawing, which is copied in text-
fig. 29, p. 104. He draws attention to the suture (s) separating
the two bones seen in section, and he states that # is the frontal
boneand y the parietal. He arrives at the conclusion that whilst
the lateral horns of the Giraffe are seen thus to originate as do the
horns of all other Pecora, in connection with the frontal bone,
104 PROF. E. RAY LANKESTER ON THE [Feb. 5,
yet that in the Giraffe the growing horn must spread from its
original position, and in fact take up a new position on the
parietal, with which he recognises that it is largely in contact
in adult life.
Text-fig. 28.
é N
y 6 \
i 2 RS
if ‘ LN
Bes ESN
ea oo" 7). 088<
Text-fig. 28.—Sagittal section through the bony tissue of the ossicone and the roof
of the skull of a very young Giraffe (same specimen as that drawn in text-
fig. 24). Drawn of the natural size.
p.oss., the ossicone; s., the parieto-frontal suture; a:., the parietal bone;
y., the frontal bone.
Text-fig. 29.—Copy of the drawing (natural size) of a sagittal section through the
ossicone and the roof of the skull of a newly-born Giraffe, published by_the late
Sir Richard Owen in the Trans. Zool. Soc. 1840.
p.oss., the ossicone; s., the parieto-frontal suture; «., stated to be a “ frontal” by
Owen but actually parietal; y., stated to be “ parietal”? by Owen but now shown
to be frontal.
_ I felt considerable doubts as to the correctness of this observa-
tion, and obtained through the kindness of Prof. Stewart some
190G3} HORNS OF THE GIRAFFE. 105
five years ago the actual skull of the newly-born Giraffe examined
by Owen, and still preserved at the Royal College of Surgeons.
The whole of the frontal and parietal regions had been cut away
from the skull and the pieces could not be found. There was
no evidence to be obtained from the specimen as to what really
was the nature of the bone a (in text-fig. 29) and the bone y. I
formed the hypothesis that Owen had had the horn-bearing region
cut out and a section made by an assistant. The section is that
which he figured and is here copied. But the piece having been
detached from the rest of the skull, Owen seems to have mistaken
right for left and back for front, so that in reality the bone
marked a is the parietal and the bone marked y is the frontal ;
and the young horn or ossicone is resting on the parietal as it
does in the later stages of growth, and not on the frontal as
supposed by Owen.
I could not test the truth of this hypothesis without examining
myself a newly-born Giraffe or a well-advanced fcetus, and accord-
ingly I have made efforts to obtain such a specimen by application
to the officials of the late African Department of the Foreign
Office and to those of the Colonial Office, as well as to naturalists
and sportsmen. Nonewly-born or feetal Giraffe came to hand, nor
could I hear of one as being preserved in any Museum in Hurope.
Accordingly I was very grateful when last summer our Secretary
was able to place at my disposal the fcetal Giraffe which was
removed from its mother after her death in the Gardens in April
1906. This feetal Giraffe was figured and described in a general
way by Mr. Beddard (Proce. Zool. Soc. 1906, p. 626), but the
examination of the skull was kindly left to me. The dead
mother of this foetus was a South-African (Transvaal) Giraffe
(G. camelopardalis wardv) and the father a Kordofan specimen
(G. camelopardalis antiquorum). Mr. Beddard has given the
dimensions of the foetus, and has estimated that it had probably
completed two-thirds of its fcetal life. He has pointed out and
figured the large size of the incipient lateral horns, and their
free extremities tufted with long hair, and has noted that their
substance 1s of a gristle-like consistency.
Soon after I received the specimen, the integument (C) was
reflected from the right side of the fronto-parietal region of the
head, under my supervision, by my assistant Dr. Ridewood ; and
by subsequent reflection of part of the periosteum the view
obtained which is given in text-figure 30. The integument of
the right side of the head (C) was thrown back, and the periosteum
of the parietal bone was reflected (D) excepting that part lying
beneath and forming the base of the right lateral horn. ‘This was
pinned down and cut away from the rest of the periosteum, leaving
it as an oval area A, marking exactly the position of the ossicone
(fibrous and soft) on the parietal bone. In the drawing the suture
separating the frontal from the parietal bone is seen (szé.), and it
is demonstrated that the base of the young lateral horn or ossicone
is wholly within the area of the parietal bone, to the periosteum
of which it is loosely attached by connective tissue.
[Feb. 5,
PROF. E. RAY LANKESTER ON ‘THE
106
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1907. ] HORNS OF THE GIRAFFE. 107
The substance of the young ossicone was free from osseous
deposit but of a tough consistence. Microscopic sections showed
it to consist of a fibro-trabecular tissue with abundant interspersed
cellular elements.
The dissection as here presented definitely establishes the fact
that the site of origin of the lateral horns or ossicones of the
Giraffe is entirely within the area of the parietal bone. The
supposition that Sir Richard Owen had by inadvertence reversed
the cut-out portion of the fronto-parietal area of his newly-born
Giraffe, and had thus identified parietal as frontal and frontal as
parietal, is confirmed. The importance of the distinction between
the genus Giraffa and the genus Okapia, arising from the parietal
position of the lateral ossicones in the former and their frontal
position in the latter, is thus placed on a firm basis, since it 1s
shown that at a stage of growth even earlier than that of birth
the “forecast” or rudiment of the Giraffe’s lateral bony cone
(ossicone) is placed and attached absolutely and solely within the
area of the parietal bone.
Text-fig. 31.
* i l) \
LZ i [Yf ye
The left “ horn ” of the fcetal Giraffe, drawn of the natural size.
A. Seen from behind (postero-external face).
B. Seen from in front (antero-internal face).
The form of the soft unossified forecasts of the lateral ossi-
cones in this foetal Giraffe is worthy of further notice. As shown
in Plate V. accompanying a later paper in this volume, and in the
text-figure here given (fig. 31), the upgrowth is of considerable size,
is compressed so as to give a narrow oblong area in transverse
section, and is set on the head so that the elongated basal area
has an oblique position, with its long diameter directed back-
wards and inwards towards the median line. The surface of the
ossicone-forecast is covered with hair, which 1s very coarse and
long at the free upstanding margin, as shown in text-fig. 31.
108 PROF, E. RAY LANKESTER ON THE [ Feb. 5,
The text-figure 32 also shows that three of the dark hair-bands
are disposed around the base of the young ossicone. The great
eZ
EA
EZ
Right Horn
base
&
RIGHT.
Diagram to show tbe flattened plate-like form and the orientation of
the horns of the foetal Giraffe.
a, b, c, d. The four left-side inter-cornual colour-bands of the pelage.
lateral compression or flattening of the young structure is
remarkable as compared with the fuller circular contour of a
transverse section after ossification has advanced.
Possible Relations of the Giraffid Ossicones to the bony Horn-
cores of Bovide and the Antlers of Cervide.
In my memoir describing Sir Harry Johnston’s specimens of
Okapi, I ventured on some speculations as to the relationships of
the bony growths called horns and antlers in the Pecora. These
speculations were vitiated by the uncertainty of existing know-
ledge as to the actual embryological origin of the structures
compared. They assumed the origin of the horn-cores of Bovidee
as separate rudiments which become united to the osteogenetic
tissue of the frontal bone at an early period of development.
Although convincing histological accounts of their development
are not yet in existence, it seems to result from the observations
of Dirst that the horn-cores of Bovide are not of independent
origin, but are actual outgrowths of the osteogenetic tissue of the
frontal bone. The same origin appears to hold for the horn-style
or column which supports the antler of the Cervide.
It is unfortunately the fact that our knowledge of the early
embryological history of the ‘ossicones” of the Giraffe and
Okapi is still more defective. All we know is (1) that in Owen’s
newly-born Giraffe the rudimentary unossified ossicone was sepa-
rated (as shown in text-fig. 29) by the dense periosteal membrane
from the subjacent parietal bone; (2) that in the feetal specimen
of two-thirds time here described the soft forecast of the ossicone
was equally cut off from the subjacent parietal bone by dense
1907. ] HORNS OF THE GIRAFFE. 109
periosteum ; and (3) that in a much younger dried skull there is
no indication on the cranial wall of any “ outgrowth.”
We shall not be ina position to speak definitely until other
foetal Giraffes of younger stages have been examined by proper
histological methods; but it seems legitimate to consider the
lateral ossicones of the Giraffe, and therefore of the Okapi, as
originating in a fibrous osteogenetic mass which gives rise to a
protrusion of the integument and originates in the connective
tissue of the integument rather than in the osteogenetic tissue of
the cranial roof, from which it is separated by a dense membranous
periosteum.
The lateral ossicones of the Giraffe appear to have the same
nature and mode of origin as has the centrally-placed ossicone of
the same animal. This median ossicone is variable in size, and
does not appear until many years after birth when growth is
nearly complete. The histological processes by which the median
ossicone is formed have never yet been studied, but it is practically
certain that it forms not as an outgrowth of the bone of the
cranial roof, but as a “dermal” or tegumentary growth external
to and independent of the cranial bone.
The same process which leads in the Giraffe to the formation of
a median ossicone, In some cases becomes specially active and leads
to the formation of additional “ supernumerary ” sub-tegumental
ossifications. Thus two such of smaller size than the normal
median ossicone are seen in the median line in a skull of Giraffe
in the British Museum, in which bony deposits on the margin
of the orbit are also seen. The skull of Okapi brought by
Capt. Boyd Alexander from the Welle River shows such an
exceptional sub-tegumental bony deposit on the margin of each
orbit ; and it is to be noticed that both in Giraffes and Okapis in
old individuals the base of each ossicone spreads very widely as a
thin encrusting layer, so as to involve much of the frontal in
specimens of Giraffe and a large surface of the parietal in the case
of Okapi.
These superficial ossicones and their outspreading marginal
growths of the Giraftide cannot be accurately marked off in later
life, although they can be separated in earlier life, from the highly
important upgrowths or “tumescences” (as I previously called
them) of the cranial bones over which they lie. The completed
lateral horn of the Giraffe consists very largely of a conical up-
growth of the parietal bone, and also of the frontal bone—occupied
by an air-sinus—developed between the tabule of the bone. The
basal region of the upgrowth involves the frontal bone more
largely than the parietal. The originally independent ossicone
contributes but a small amount to the whole bulk of the structure.
It forms merely the terminal knob, and is fitted over the tumes-
cence like a superficial investment which dwindles in thickness as
it descends the cone until it becomes a mere film.
The position of the lateral and median ossicones in an adult
110 PROF. E. RAY LANKESTER ON THE [Feb. 5,
(though not senile) Giraffe, and the relation of the sinus of the
parietal and frontal to the lateral horn, is shown in the figures (text-
figs. 33 & 34) which were prepared from photographs of sections
of a skull made under my direction. It will be observed that it
is not possible in such sections to distinguish the line of ankylosis
of the separate bones; we can only guess somewhat vaguely as to
what belongs to each of the three elements fused together, viz.,
ossicone, parietal, and frontal.
Text-fig. 33.
Skull of Giraffe, left side; key-figure, a little larger than one-sixth (linear) of the
natural size, showing the directions in which the skull was cut.
(From the Proc. Zool. Soc. London, 1904, vol. i. p. 151.)
Thus we see that the “‘ossicone” of the adult Giraffe is essen-
tially but a cap of bony substance fitting over the great upgrowth
or conical tumescence of the cranial wall—at the first most marked
in the parietal area,—and that it makes its first appearance as a
solid growth of fibrous tissue resting on the flat cranial roof.
The same conclusion may legitimately be drawn from what we
can see of the early and later conditions of the ossicone in
Okapi*.
* The tumescence of the frontal bone of each side in Okapi, which ankyloses with
the ossicone overlying it, is of considerable volume, like those to which the parietal
gives rise in the Giraffe. I shall have an opportunity of describing this structure
more fully hereafter. At the present moment I desire to draw attention to a curious
fact with regard to the rudiment of a median horn in Okapi. The base of the nasal
bone gives rise to a small but well-marked median tumescence in the Okapi. I
described this in Sir Harry Johnston’s specimen (Trans. Zool. Soc. vol. xvi.). In horn-
bearing skulls of Okapi this is more pronounced than in the hornless speeimens ; it
1907. | HORNS OF THE GIRAFFE. Wail
Since it is clear then that the Giraffe’s horns comprise two
bony factors, the question arises which of the two corresponds
with the horn-core of Bovide, or whether either or neither had
such a correspondence.
Text-fig. 34.
we3)s, parrect
\ horn
2 ul SGM
VAY
alisph ; “y Vi ‘
7 |
bus sph . pteryg.
View, looking forwards, of the skull cut in the direction of the line 4 in text-fig. 33.
One-third (linear) of the natural size.
(From the Proc. Zool. Soc. London, 1904, vol. i. p. 154.)
is more strongly marked in the horn-bearing skull of Okapi brought by Capt. Boyd
Alexander from the Welle River than in any other specimen seen by me. No “ossi-
cone” or tegumentary cap has been observed in Okapi in connection with this
median tumescence. Has there been one present in the ancestors of Okapi? Or
does the tumescence precede the formation of a tegumentary ossicone ?
112 PROF. E. RAY LANKESTER ON THE | Feb. 5,
Text-fig. 35.
i] te £
Bi) 4 Oi
Wy; ip Chis ae
MM 27
KOEN 7 /
Drawing, of the natural size, of a young foetal skull of a. Giraffe, preserved in
the British Museum. ‘he foetus is at so early a stage that no trace of the
future horn or ossicone is presented, and moreover the arietal bone does not
form the vertex of the brain-case as in the skull drawn in text-fig. 24, p. 101. It is
worthy of remark that a line drawn from the occ.-par. suture to the fronto-par.
suture is in this early foetal skull almost at right angles to a line drawn from
the latter to the naso-frontal suture, a condition which is even more strikingly
exhibited in the adult skull of the Common Ox (Bos) as contrasted with that of
Ovis, Ovibos, and Antilopide. In adult Giraffe the surfaces of the frontal and
parietal are more nearly parallel, and in Okapia quite so. It is a very curious
fact that in Bos the whole posterior region of the brain-case formed by the
parietals, which in Okapia stretches horizontally backwards from the fronto-
parietal suture to a distance as great as that occupied by the frontal portion, is
abolished! The parietals are vertical and not horizontal, and a sort of false
occipital ridge is formed by the fronto-parietal suture. The very young Giraffe-
foetus approaches this condition.
Tn answer to this question, it appears to me that the following
statements are justified :—
1. Since the bony horn-core of Bovide originates as a part of
the osteogenetic tissue of the frontal bone, it cannot
(according to our present knowledge) be identified without
considerable qualification with the free tegumentary t
ossicones of Giraffide.
2. The upgrowth (tumescence) of the frontal bone in Okapi and
of the parietal and frontal in Giraffe, which forms the
bulk of the lateral horns in those animals, is more nearly
similar in nature to the bony horn-cores of Bovide than
are the ossicones of Girattidee.
3. The frontal tumescence or upgrowth of Okapi cannot be
considered as morphologically identical with the parietal
tumescence or upgrowth of Giraffa.
4. The free lateral ossicone of Giraffa might legitimately be con-
sidered as morphologically identical with the free lateral
ossicone of Okapi. The free tegumentary ossicone of an
1907. ] HORNS OF THE GIRAFFE. 113
ancestral form might in a series of generations shift its
position from the frontal to the parietal area ; and it might
reasonably be admitted that the upgrowth or tumescence
of the frontal ceased to develop when the parietal position
was assumed by the ossicone with consequent tumescence
and upgrowth of the parietal bone.
5. On the other hand, the theoretical assumption that the
frontally-placed ossicone of Okapi and the parietally-placed
ossicone of Giraffe are independent of one another and
possibly co-exist in an ancestral form, is favoured by the
fact that the Giraffe does develop a third well-marked
ossicone in the mid-line of the frontal, and that both
Okapi and Giraffe exhibit minute supernumerary growths
of the kind on the cranial surface.
6. It results from these considerations that it is not possible at
present to trace the lateral horns of the Okapi and the
Giraffe into any close genetic relationship with those of
Bovidee—still less of Cervide. At the same time it is
possible that the peculiar superficial element of the bony
horn (the ossicone) is identical in the lateral horns of
Okapi and Giraffa, having shifted its position backwards
in the latter genus. This conclusion is not, however, by
any means forced upon us since the Giraffide are known
to have an additional ossicone—the median one; and it is
therefore not without analogy that independent frontal
and parietal G sicones should develop.
I am aware that it is not difficult to make assumptions by
means of which a genetic relationship between the lateral horns
of Giraffide, Bovide, and Cervide is rendered possible; but it
should, I think, be clearly understood that there is at present no
direct evidence to support these assumptions. It may be assumed
(a) that a bony horn of the nature of the horn-core of the Bovide,
or of the antler-column of Cervide, has in some remote ancestors
of the Giraffidee become segregated from the frontal bone of which
it was a part, and acquired independent existence as a fibrous
rudiment as well as independent ossification, thus establishing the
independent lateral ossicone of the Giraffide. Or, again, it may
be assumed (6) that in ancestors of the Bovide and Cervide,
bony horns which were existing as free tegumentary products,
ankylosing in mature age with subjacent cranial bones, became
so ankylosed at earlier and earlier stages of development until all
trace of their independent origin was lost, and they appeared to
originate as growths of the frontal bone itself. The stock so
endowed gave rise (it would be assumed) to Bovide and Cervide ;
that portion of the ancestry which retained the original method
of development of free tegumentary ossicones became, on the other
hand, the progenitors of the Giraftide.
I am not aware of any facts in the structure of living or extinct
Artiodactyla which furnish an analogy for either of these pro-
cesses of transformation. Nor do I think that our knowledge of the
extinct forms such as Samotheriuwm, Helladotherium, Sivatherium,
Proc. Zoou. Soc.—1907, No. VIII. 8
114 ON THE HORNS OF THE GIRAFFE. [ Feb. 5,
Bramatherium, and Hydaspitherium, is of a nature to assist in
establishing the existence of such a remarkable transformation.
The assumption a—namely, that the transition was from frontal
outgrowths such as Bovide and Cervide present, to the segregation
and independence of the Giraffid ossicone—meets (it may be
pointed out) with an almost insuperable obstacle in the free median
frontal ossicone of the Giraffe, for which there is no forerunner
in the Bovi-Cervine scheme of solid continuous outgrowths of
the eranial wall. The consideration of the Lower Miocene Proto-
ceros with its four pairs of horns does not help us much in this
question, though its possession of a pair of parietal and of two
pairs of frontal upgrowths or ‘ bosses’ is significant.
Various Positions occupied by the Paired Frontal Horns of
Cavicorn and Cervine Ruminants.
An interesting fact in relation to the question of the identifi-
cation of Giraffe’s parietal with Okapi’s frontal ossicones is that
Text-fig. 36.
-—-NASAL BONE.
Anterior horns
FRONTAL BONE.~ ~_
oy Be of Tetraceros.
= a
Antilocapra americana ~
Qreotragus saltator.
__Ovibos moschatus,
Cervus elaphus.
Ovis tragelaphus __ _ _
Nemorrhedus goral. _ Posterior horns
of Tetraceros.
“Alcelaphus caama.
Cephalophus coronatus.
PARIETAL BONE.
Diagram to show the various positions on the frontal bone at which the bony horn-
cores of the Cavicorn and Cervine ruminants may take their growth. Though
ranging widely over the frontals these upgrowths are never found as part of the
parietal bones.
PW
ISO
Paes:
SUED) Ol AA IWOETAIL, CiRAR FIs.
( Half the natural size.)
1907.] | ON HAIR-FRINGES ETC. ON THE FACE OF GIRAFFES. 115
the paired frontal horns of Antelopes, Bovines, and Deer occupy
very widely-separated positions in different genera. Dr. Ridewood
has kindly prepared for me a diagram (text-fig. 36) showing these
varied positions in a series of genera. The most remarkable
position is that of the horn-cores of the Antelope Cephalophus,
which. is not fully exhibited in the diagram. The frontal in
some species of this genus actually pushes out a process into the
area of the parietal, upon which the horn-core rests. The horn-
cores are seen (by reference to the diagram) to arise sometimes at
the hinder margin of the frontal, sometimes on the orbital ring,
sometimes in the anterior third of the frontal. The co-existence
of two pairs in Tetraceros suggests a multiplicity of horns in
ancestral forms.
The position of the horn-cores in the genus Bos is not marked
in the diagram ; it is identical with that of the posterior pair of
Tetraceros.
2. Parallel Hair-fringes and Colour-striping on the Face of
Foetal and Adult Giraffes. By HE. Ray LANKEsTER,
VeAn Osc. tO. RIS. EZ.S., Direction ot the
British Museum (Natural History).
[Received February 5, 1907. ]
(Plate V.* and Text-figures 37-48.)
When examining the feetal Giraffe which I received from the
Society in the past summer, I observed a number of parallel
bands or stripes of dark and light colour on the hairy coat of the
Text-fig. 37.
Left side of the head of the foetal Giraffe described, showing colour-stripes on the
snout and above and below the eye. The small arrows indicate the direction of
slope of the hairs. About three-tenths of the natural size.
* For explanation of the Plate, see p, 125.
g*
116 PROF. E. RAY LANKESTER ON HAIR-FRINGES AND [ Feb. 5,
face—between the two lateral horn-sacs, also between the eyes and
above the eyes, which are carefully represented in the coloured
drawing (Plate V.) of the face of the foetal Giraffe. I observed
sinilar but more strongly marked and broader bands of alternating
reddish-brown and paler colour between the nostrils (see Plate V.
and also text-fig. 30, swpra, p. 106) and at the side of the upper
lip, on the front of the lower jaw and below the eye (text-fig. 37,
p. 115).
rae colour-bands between the horns and the eyes varied in
intensity according to the angle of the incident light, and could
be temporarily destroyed by pressing the skin and smoothing
down the hair. After careful examination of the hairs, I came
to the conclusion that there was no actual difference of colour in
the hairs occupying the darker stripes and those placed on the
lighter tracts, but that the phenomenon was due to the existence
of parallel linear depressions or wrinkles the existence of which
was made obvious by a transverse section of the integument
(text-fig. 38). The hairs are crowded together in the trough of
the wrinkle, and further apart in the convex intermediate areas.
That “ wrinkling” could produce such an impression of dark and
light banding was demonstrated by the casual folds and wrinkles
of the integument on the legs, and by purposely producing such
wrinkles by pressing or folding the hairy integument.
Text-fig. 38.
Sister a = =
Su oy a eee 2
yp = — Ge = =
kee (ASS ae eS SS ea
ws Do ES SE 8p SS SS Se
ee NS SE Se ==
= SSS
Section across three dark-coloured bands above the eye of the foetal Giraffe, showing
three longitudinal furrows or depressions in transverse section corresponding to
the dark bands. Magnified.
Nevertheless I could not attribute the colour-banding to a
mere accidental or casual formation of wrinkles. Their definite
form and arrangement precludes such an explanation. They
appeared to me to be the expression of a definite structural
condition. Moreover, immediately over the eyes and on the
snout the difference of colour of the alternating bands was very
strong, the darkly-coloured stripes being of a strong reddish-
brown tint and the intermediate bands quite pale; and in this
case a pigment was present in the hairs of the darker stripes
which was not so richly developed in those of the lighter
neighbouring stripes. I found, on microscopic examination of
the hairs, that they could be roughly divided into three sizes ;
1907. ] COLOUR-STRIPING ON THE FACE OF GIRAFFES. ugly,
the largest, few in number ; medium-sized, more numerous; and
a smallest size, the most abundant (text-fig. 39). The two larger
sizes of hair were remarkable for appearing almost colourless in
their lower (proximal) moiety and very dark in the upper (distal)
Text-fig. 39.
[Be iB 7
HAA FF
Section of the skin of the frontal region of the foetal Giraffe, showing
three sizes of hairs. Greatly magnified.
moiety. The smallest hairs were dark-coloured throughout.
When a piece of the hairy colour-banded region was examined
with a strong lens, the tips of the hairs were seen to converge from
the two sides of the shallow furrows or wrinkles as shown in the
118 PROF. E, RAY LANKESTER ON HAIR-FRINGES AND | Feb. 5,
text-figure 40. The superposition of the darker free ends of
the hairs conceals theix paler basal regions, and thus intensifies
the difference between the apparent colouring of the troughs or
wrinkles and the intermediate spaces where the hairs do not
converge. An interesting experiment in regard to this matter was
made by my assistant Dr. Ridewood. He took a piece of pale
hat-maker’s plush, and stained with dark pigment the free ends
of its hairy surface. He found that on throwing this manufac-
tured material into a series of wrinkles, very strong alternation
Text-fig. 40.
SS
SSS
SS —
—— SS
—=
=
Surface-view of banded “pelage”’ of the frontal region of foetal Giraffe. showing the
convergence of the hairs at the three longitudinal bands of dark appearance.
Magnified.
of dark and light colour-stripes could be produced. (This pre-
pared material was shown to the Meeting and dark and light
bands produced in it and removed at pleasure by alternately
throwing it into wrinkles and stretching it so as to remove the
wrinkles.)
Although these colour-stripes on the head of the fcetal Giraffe
thus appear not to be due to alternate tracts of hair of differing
colour, it seems to me that they have a real existence as effective
colour-marking, and that their structural cause is to be found in
the differentiation of the attachment of the panniculus carnosus
1907. | COLOUR-STRIPING ON THE FACE OF GIRAFFES. 119
Text-fig, 41.
An enlarged drawing of a single hair of average size from the fr
ontal region of an
adult Giraffe (Giraffa camelopardalis wardi), showing the dark pigmented
free extremity and the opaque white lower portion.
120 PROF. E. RAY LANKESTER ON HAIR-FRINGES AND [| Feb. 5,
(ih a
_— a 1
aah Za
= £m
Za Zi
= = F
S—
Ny i i iN
ww
Colour-striping on the face of an adult male Giraffe from Kordofan (G. c. antiquorum).
rawn from a specimen living in the Society’s Gardens.
Wes Text-fig. 43.
Z
A nN *&
ZY
TT i:
BZ o
B oI
ZB i "
a co |
th Ay NG &
(i
re
Colour-striping on the face of an adult female Giraffe from Kordofan (Gc. antiquorwm).
rawn trom a specimen living in the Society’s Gardens.
1907.] COLOUR-STRIPING ON THE FACE OF GIRAFFES. 121
to the integument, and in the related direction and convergence
of the hairs in definite bands or hair streams. An examination
of the adult Giraffe establishes the truth of this view. I visited
the Giraffe-house at the Society’s Gardens in order to examine
the father of the feetal specimen in which I had observed this
colour-striping. The father is a Kordofan specimen, and there
is also in the Giraffe-house a female Kordofan Giraffe. I was
not a little surprised to find very strongly-marked colour-striping
over the eyes of both these Giraffes, especially well-marked in
the female. Portions of the muzzle are also banded, and between
the eye and the angle of the mouth are developed from four to six
Text-fig. 44.
Nay
SS
S GN
ie & |
Set il
Legh
Bat ae
y
|
AE
\\
\
Colour-striping on the face of Giraffa camelopardalis cottoni from Mt. Elgon,
Uganda. This specimen shows a horn-like exostosis over the right eye. Drawn
from a specimen in the British Museum.
strongly-marked horizontal colour-bands which are not present in
the foetus (see text-figs. 42 & 43). Ifound that when the eye was
shut and the skin above the eye stretched, the strongly-marked
dark and light bands disappeared giving place to an irregularly
blotched appearance *, which immediately resolved itself into
alternate dark and light bands when the eye was opened and the
* It seemed to me that there were in this region definite tracts in which the
pigmentation of the hair was pale, and others m which it was dark, but exact
observation was, I found, impossible in the living animal.
122 PROF. E. RAY LANKESTER ON HAIR-FRINGES AND [Feb. 5,
superciliary region thrown into the normal condition assumed
when the animal is alert. I also noticed that though the
horizontal bands between eye and mouth never actually disappear,
they are intensified by a muscular contraction resembling a
sneer which sometimes is exhibited by the Giraffe. I found that
the third Giraffe in the Gardens (a West African specimen) did
not exhibit any colour-bands on the face.
Text-fig. 45.
Colour-striping of the face and muzzle of a Transvaal Giraffe (Giraffa camelopar-
dalis wardi). Drawn from a specimen in the British Museum.
I next proceeded to examine from this point of view the
fine series of Giraffe heads and necks exhibited in the public,
gallery of the British Museum, as well as several unmounted
skins. I found that several of these specimens exhibit colour-
banding over the eye and some of them below it, whilst these same
specimens exhibit strong horizontal “fringes” or banding of the
hair between the eye and muzzle, and some also show banding on
the lower lip (see text-figs. 44-48). On the other hand, some of
the specimens exhibit little or no trace of these bands. In none
have I found any trace of the bands in the mediad position
between the horns and between the eyes, shown by the feetal
Specimen drawn in Plate V., which itself shows no trace of the
pre-orbital horizontal stripes.
1907.], COLOUR-STRIPING OF THE FACE OF GIRAFFES. 123
The bands of alternate light and dark colour are as much
larger and wider in the adult Giraffe as are all its dimensions
larger than those of the fetus. But a fact in regard to the
banded appearance of the hairy coat of the face has come to light
in the case of the adult skins which is indicated in the enlarged
drawing of a piece of the foetal pelage (text-fig. 40, p. 118). This
is that the bands are essentially due (at any rate, those which are
peculiar to the adult and most strongly-marked, viz., the hori-
zontal pre-orbital group of bands) to a differentiation of the hair
into parallel tracts of more Coase placed hairs, the points of
Text-fig. 46.
WX
cil
Wy, co
AY WHI HIN\\, \\ WAY Ay
NAIK
‘
Ml ea
\
TAK Ain
yt UG We
(eee wt UY,
SHAN
nN SS
h SW
KUNNEN
cc AS iN \ ND Wy
att vif) Wy
\) \ Wie Wy
Y,
if Mil
{i
ae
iy)
is
ee Hf Uf
if} f Uji
a Huh thi yh Hy
The same specimen as that shown in soars 45. in this drawing the colour-effect
is ignored, and only the ridge-like arrangement of the hair on the face and
muzzle in parallel fringes is shown.
which converge and stand up so as to form a well-marked raised
stripe, “fringe” or “ridge” *, and intermediate tracts of smooth
flat-lying hair. In some of the Museum specimens the more
crowded upstanding hairs appear to be coloured more darkly than
those of the intermediate tracts, but really the colour-effect is
due to the pigment of the free ends of the hairs showing, whilst
the thicker cortical substance of the bases of the hairs is white
* The ridge of upstanding hair corresponds, it must be noted, to what isa shallow
wrinkle-like groove in the feetal skin.
124 PROF. B. RAY LANKESTER ON HAIR-FRINGES AND [Feb. 9,
wth .
pie aN
rR oe PP.
th co pe
an 4 ae 4
iif INN
Wn PEN
Colour-striping (very slightly developed between eye and nostril) of the face of a
Somali-land Giraffe (Giraffa reticulatus). Drawn from a specimen in the
British Museum.
(ile, Moron
WYN
Z) al
io
2
( yp ) AK x
Ve
La, Le Sit Ha
ve
7 &. \ nc
y
\ “i WY
Hie iy
/ Yi; ty
Wy Yh
YMG TON
TATE iif YY Yi if fh
a Wi) Hii M/1, HT Wg Lyd) Hi
Hi Hi Ha hil fii Ui Wy Hd yi'f WK, iy
Walt) HI Wh WH. iol
The same specimen as that shown in text-fig. 47, but with all colour omitted. The
parallel bands between eye and nostril are not marked out in colour, but are
merely elevated fringes or ridges of hair traversing a colourless region.
1907.] COLOUR-STRIPING OF THE FACE OF GIRAFFES. 125
(text-fig. 41, p.119). In one Somali specimen (@. reticulatus) the
hairs are entirely white over a large part of the region of the face
where the horizontal bands are developed. Yet these horizontal
bands show up very distinctly on account of the more erect setting
of the hairs (see text-figs. 47 & 48). In some cases the hairs of
these ridges seem to have yielded more readily to destructive
processes connected with taxidermy than have the hairs of the
neighbouring tracts, and consequently the bands are marked out
by nearly bald furrows or pathways.
Another point of interest is that one of the horizontal pre-
orbital hair-bands in the adult Giraffe directly leads up to the
pre-orbital hair-whorl, and that the position of this hair-whorl
appears to be farther in front of the eye in the Somali G. reticu-
latus than in other Giraffes, whilst undoubtedly the hair is more
erect and strongly developed on those ridges and on the hair-
whorl in that species than in the other Giraffes which I have
been able to examine.
It would be interesting to ascertain how far the varying
development of these colour-bands and parallel hair-ridges on the
face of the Giraffe is constant in the different local varieties of
Giraffe which have been distinguished. This isnot a matter with
which I am at present able to deal. The purpose of the present
communication is to call the attention of zoologists to a banded
structure of the integument of the face in Giraffes which results
in the appearance of dark and light parallel bands of hair and in
the formation of strongly-marked parallel fringes or ridges of
erect hair separated by bands of recumbent hair. These struc-
tures are recorded in a foetal and in adult Giraffes, and appear to
have hitherto escaped attention.
The production of optically effective colour-bands by the mere
crowding and direction of hairs along certain lines, without actual
difference :n the pigmentation of the hairs of the lighter and
darker stripes, has possibly some significance in regard to the
origin and development of the more definite striping of the
mammalian pelage so frequently shown, when dark and light
stripes distinguished from one another are caused by the actual
presence of pigment in the hair of the dark stripes, and its
absence in the hair of the lighter or white stripes.
EXPLANATION OF PLATE V.
Coloured drawing of the head and face, seen from above, of the foetus of a
Giraffe removed from its mother which died in the Society’s menagerie in April
1906.
The head of the foetus measured seven inches and one quarter from the anterior
angle of the base of the outer ear to the extreme border of the upper lips.
The specimen had been preserved in alcohol for six months.
Details concerning the parentage &c. of this foetus are given by Mr. Beddard in
Proc. Zool. Soc. 1906, p. 626.
126 PROF, E, RAY LANKESTER ON [ Feb. 5,
3. On the Hxistence of Rudimentary Antlers in the Okapi.
By E. Ray Lanxester, M.A., D.Sc., LL.D., F.B.S.,
F.Z.8., Director of the British Museum (Natural His-
tory).
| Received February 5, 1907. |
(Plates VI. & VII.* and Text-figures 49-55.)
We know a great deal more as to the horns of the Okapi than
was the case when I communicated my description of that animal
to the Society in 1901, and founded the genus Okapia.
The two skulls sent home by Sir Harry Johnston—the first
seen in Europe—were hornless, and it was at first a matter of
doubt as to whether the Okapi wasa hornless Giraffid, or whether
the male possessed horns whilst these two skulls were the one
immature and the other that of a hornless female.
During the printing of my memoir additional specimens were
received in Brussels, and were transmitted to Dr. Forsyth
Major in London for study and description. I saw in Dr. Forsyth
Major’s possession a fine adult Okapi skull which had a pair of
well-developed bony cones rising each by a broad base from the
frontals, of which they appeared to form part. No suture was
visible. An outline of this skull was published in my memoir
by kind permission of Dr. Forsyth Major. I also was able to
examine and to mention the existence of a curious structure
discovered by that gentleman in regard to these ossicusps ; and I
described it in the following terms :—‘“‘ The fine bony cones
three inches long, which have made their appearance in the
Text-fig. 49.
Drawing of a fore and aft section through the tip of the ossicone of an adult Okapi
in the collection of the Museum of the Congo Independent State. The section
and drawing were made by Dr. Forsyth Major.
The section shows the penetration of transverse fissures from the surface into the
interior of the horn-tip.
a, dense ivory-like bone; 6, posteriorly-placed transverse fissure; c, more anterior
transverse fissure (marking off and presumably about to cut off and detach an
anterior seement or plate of bone as a rudimentary “ antler ”)
Brussels skull, show no suture at their base, nor any indication
of origin as separate cap-like structures. For all that one can see
they may be direct outgrowths of the frontal bone itself. Curiously
* For explanation of the Plates, see p. 134.
ici LON eae anvils
Left. | Right.
Left. Right.
4
H.Grénvold dei London Stereoscopic Co imp-
PLOT N= ene S Ol Oi i
eas LOO, see valle
G.M Woodward del. London Stereoscopic Co imp.
YOUNG OSSIGONE OR HORN OF-OKAPI.
ie Pal
it Tne
TRC ALTA ts: ue Als
‘ 4, a pert v
ow her) by
1907. | RUDIMENTARY ANTLERS IN THE OKAPI. 127
enough, the point and posterior margin of the bony cone are
polished as though it had protruded through the skin like a
cervine antler. The point is separated by a suture from the rest
of the ‘ ossicone,’ forming a small terminal cap of bone a third of
an inch in depth. This curious structure, as well as a possible
second suture a little lower down the ossicone, was pointed out to
me by Dr. Forsyth Major. These appearances will be figured in
that gentleman’s memoir on the Brussels’ specimens.” This is
the first and so far the only published notice of the antler-like
tips of the Okapi’s horns. The figure prepared by Dr. Major of
the section made by him through the end of this Okapi’s ossicone
is reproduced in the text-figure here appended (text-fig. 49).
Dr. Major does not himself propose to publish anything further
at present on the Okapi, and the little drawing has been placed
in my hands by him. A tracing of it was also kindly sent to me
by M. Fraipont, of Liége.
The further history of our knowledge of the horns of the
Okapi has been complicated by the arrival in Europe of various
specimens, concerning the sex of which either erroneous informa-
tion or none at all has been given by the natives from whom the
specimens were obtained. Thus Dr. Forsyth Major was led to
suppose that the female Okapi has a small unattached ossicone,
some two inches in length, when adult, but he subsequently came
to the conclusion that this supposed female was in reality a young
male. In ‘ La Belgique Coloniale,’ No. 21, May 1902, Dr. Forsyth
Major wrote :—“ L’Okapi posséde deux cornes frontales, recouvertes
dune peau velue, plus petites, de forme conique et presque verti-
cales chez la femelle; plus grandes, dirigées obliquement en
arriére et en peu triangulaires chez le male.”
At a subsequent date Dr. Forsyth Major came to the conclu-
sion that the specimen supposed to be a female possessing small
j * ossicones, was in reality a young male (Proc. Zool. Soe.
». 339), and that the female Okapi is hornless, whilst the
ie possesses “horns” which make their appearance as
conical structures, ossifying independently of the sub-
nes (as in the Giraffe) and becoming firmly ankylosed
mtal bone in the adult—a boss-like upgrowth of which
0 the structure of the complete horn.
s little room for doubt that this is the true account of
r, though we still are in want of full information as to
icters of the adult female Okapi*. In a subsequent
vation I shall be able to give more precisely the characters
» types of skull, supposed to be that of the horn-bearing
the hornless female, respectively. The skulls carrying
sed or merely loosely-attached bony cone on the frontals
‘0 the uncertainty which exists as to the origin of skin and skull which
»s sent home from Africa as belonging to one individual, whereas they
in certain cases belong to distinct individuals, it is still doubtful as to
wuetner tne female Okapi has or has not in the adult condition a small knob-like
protuberance of the integument, separable from the subjacent bone and representing
the horn of the male.
128 PROF, E. RAY LANKESTER ON [ Feb. 5,
(supposed to be those of males) are longer and narrower than the
equally large or larger skulls devoid of any bony cones in connec-
tion with the frontals (supposed to be those of females) *. What-
ever opinion is held, or whatever decision may be ultimately
arrived at in regard to these two types of skull, it is the fact that
they are very distinct from one another and that all the Okapi skulls
which I have examined can be definitely assigned to one or the
Text-fig. 50.
a.
Rudimentary free ossicone of hemispherical shape from the skin overlying the frontal
bossed region of the skull of an Okapi of the broad-skulled type—sub-adult
(deciduous molars very much worn, premolars not yet visible ; third lower molar
in use on both sides, fifth cusp shows slight wear).
a, natural size; 6, enlarged.
Text-fig. 51.
Section of the ossicusp drawn in text-fig. 50, to show the incomplete ossification.
other of these two types. There is no third form known. The
two types may perhaps be best distinguished as O. johrestoni (the
name I gave to the broad hornless sub-adult skull accompanying
* One of these broad-skulled specimens has, however, been found to possess a pair
of completely detached bony ossicones of minute size embedded in the integument.
The specimen is a little older (as indicated by the dentition) than Sir Harry John-
ston’s larger individual (that mounted in the British Museum), but is not quite
adult. It belongs to Messrs. Rowland Ward. I give here figures of the minute
ossicone (text-figs. 50 & 51).
1907. ] RUDIMENTARY ANTLERS IN THE OKAPI. 129
the skin sent home by Sir Harry Johnston) and 0. liebrechtsi,
the name given by Dr. Forsyth Major to the more elongate and
narrow type of skull, which is that usually provided with bony
cones attached to or ankylosed with the frontal bones. It is
important to note that Dr. Major figures a skull (Proc. Zool.
Soe. loc. cit. p. 423) which is hornless and is regarded by him
as that of a female of the elongate type, O. liebrechtsi. I hope
shortly to publish some measurements and outlines of these two
types of skull. I have examined three of the UO. johnstoni-type,
and five of the O. liebrechtsi-type. Though there is considerable
variation in the number and breadth of the white stripes on the
fore and hind limbs of the skins of Okapi received in this country
(including the excellent specimens obtained independently by
Major Powell Cotton and by Captain Boyd Alexander from widely
separated localities, the former from the Ituri Forest, the latter
from the Welle River), I have seen no evidence that a different
striping of the skin is associated with the difference of skull-form.
On the contrary, there is positive evidence that the striping of
the skin is very nearly identical (though no two specimens are
exactly alike) in animals which possessed the liebrechtsi form of skull
with that exhibited by the mounted specimen (0. johnstoni) with
hornless skull, sent home by Sir Harry Johnston, figured by me,
and now in the British Museum. Nevertheless, it is true that
direct and convincing evidence is as yet wanting for the conclusion
that O. liebrechtsi is merely the male of O. johnstoni.
When I had an opportunity (in 1904) of examining the fine
skin of the adult (supposed) male Okapi, presented by the Congo
State to the Museum of Paris, which is set up in the publie
gallery there, I was especially anxious to note the state of the
horn-tips. I found that they were represented in the mounted
specimen and were seen projecting through the skin which
clothed the ‘‘ossicone” up to a limit of about half-an-inch from
the tip. From this level the dense bony matter was naked.
It showed in each horn two fine transverse grooves, as 10
the ossicone examined and sliced by Dr. Forsyth Major. This
went far to prove that the condition noted by him was not
exceptional or morbid, and accordingly I have examined the
ossicones of other specimens of adult male Okapis, as opportunity
occurred. Several skins and imperfect skeletons have been
received in London by dealers in zoological specimens, and I am
especially indebted to Messrs. Rowland Ward & Co. for the
opportunity of examining the ossicones of four adult Okapis.
Of two of these individuals I have had the ossicones drawn
(PI. VI.) so as to show the free termination from different points
of view. The two other specimens examined by me presented
the same remarkable appearances as those figured, and as shown
by the Paris Okapi, but I was unable to procure carefully drawn
figures of them. Thus, including the Brussels skull examined
by Dr. Forsyth Major, I have ascertained the existence of these
transverse grooves or fissures in six adult male Okapis. I have
Proc. Zoou. Soc.—1907, No. IX. 9
130 PROF. E. RAY LANKESTER ON [ Feb. 5,
also evidence of their existence in a plaster cast of another
specimen which passed through the hands of Messrs. Rowland
Ward & Co.
An examination of the figures given in Plate VI. shows that
in all four ossicones (the right and left of two adult male Okapis)
the free terminal region is smooth and polished, forming a cap of
about half an inch in length, whilst this region is followed by a
rougher substance, furrowed on the surface. The polished region
projects beyond the skin, the rougher region is clothed by the
living integument. In all there are very deep horizontal fissures
in the polished material of the “cap.” These fissures are some-
what irregular in form, and it is impossible without making a
section (which I had not permission to do)* through the solid
material to ascertain their depth. They are of the same nature as
those shown in the text-figure in section (text-figure 49, p. 126).
I think there can be little doubt that these transverse fissures
are caused by the ingrowth of the living tissue after the pro-
trusion of the dense polished cap, so as to cut off the protruding
portion and provide for its breaking off—Just as an antler is cut
off and prepared for disruption in the Cervidee. A small conical
piece is thus thrown off from the end of the horn or ossicusp, and
may be regarded as a rudimentary or minute “antler.”
But the process of discarding these minute points or antlers in
the Okapi appears to differ from what occurs in the Cervide, not
only in the minute size of the discarded segments, but in the fact
that the preparation for the breaking off a second (and even a
third) segment takes place before the first piece has been got rid
of. The living tissue having absorbed the bony matter by a
horizontal ingrowth and having created a transverse break in the
continuity of the osseous substance (see text-figure), recedes for a
distance of a sixth of an inch or less, and then again penetrates
inwards, forming a new horizon of disruption; and from the
appearance of the specimens figured in Plate VI., especially figs. 4
and 5, it seems that this process of the recession of the living
investment of the horn-tip and the subsequent ingrowth of the
living tissue, may be again repeated before the most anterior
piece is broken off, so that these horizontal fissures are visible on
the surface of the horn-tip, following one another somewhat
irregularly.
* Since reading this paper, I have been kindly permitted by the authorities of the
Royal Scottish Museum to examine the horns of one of the specimens above referred
to by making a section of the tip of the horn. The piece cut out has been drawn
and then carefully replaced and cemented in position, so that no injury is done to the
specimen. The skull lent to me by the Royal Scottish Museum is that of which I had
already drawn the horn-tips in figs. 1 to 8, Plate VI., before it had passed from
the possession of Messrs. Rowland Ward & Co. The sections drawn on an
enlarged scale in text-figs. 52 & 53 explain themselves. It is seen that the grooves
or fissures visible on the surface do not extend very deeply, but that there is evidence
of resorptive activity in the form of certain branching canal-like structures lying
deeply within the bony matter, which have probably been excavated by resorptive
ingrowths from the soft surface tissues.
1907. | RUDIMENTARY ANTLERS IN THE OKAPT. 131
Text-fig. 52.
Post.
A. B.
The diagram C shows the direction im which certain cuts have been made in the
left osseous horn (ossicone or ossicusp) of the Edinburgh Okapi (also illustrated
in P). VI. figs. 1 to 8).
a, anterior border; b, posterior border.’ L, left side; R, right side.
A isa drawing three times the natural size of the cut surface of the bisected horn,
the bisection being effected in a plane erected on the line ab of the diagram C.
It shows the eating in of the transverse fissures into the dense bony substance,
and a number of irregular spaces and fissures which are probably cavities due to
re-sorption of the bone. B is a drawing of the other half of the same bisected
horn-tip.
Text-fig. 53.
Drawing of three times the natural size of the surface of a section obtained by
cutting half of the same bisected horn-tip through a plane erected on the line R
of the diagram, separating the shaded from the unshaded area. The penetration
of the transverse fissures is shown. The cavities m, ”, 0 correspond to the
transverse fissures labelled m, n, 0 in fig. 5 of Plate VI., representing the same
specimen before it was divided. A
9?
132 PROF. E. RAY LANKESTER ON Feb. 5
>
We have no knowledge or indication that the shedding of these
demarcated segments of the horn-tip is seasonal, nor indeed of
the actual occurrence of such a shedding. However, the animal
which is referred to in the explanation of Plate VI. as specimen A,
shows in both right and left horn-tips a well-marked concavity,
such as would correspond to the scar left by the “‘ shedding” or
breaking off of a previously existing cap or segment of the dense
bony substance. In fig. 2, this is seen in a view of the mediad
(inner) face of the right ossicusp, and is marked w. In fig. 8 the
same cavity is seen (and marked x) in a view taken from in
front of, and somewhat above, the same horn-tip.
It seems to me that we have in these appearances of the Okapi’s
free or naked horn-tip (external termination of the ossicone) the
evidence of a process of the same physiological significance as
that seen in the seasonal removal of the antler in the Cervide.
The continued contact of a deep tissue such as bone with the
infective material of the outer world cannot be tolerated : necrotic
organisms must effect a lodgment and gradually extend their
ravages into the whole tract of the ossicone, and even to the
bones of the skull. Accordingly, the exposed “tip” is cut off by
a hone-absorbing ingrowth of the living tegumentary tissue, and
the process of autotomy is active and recurrent. In the Okapi
the process appears to be less elaborated and regularised than in
the Cervide, and we do not know at present the details of its
commencement or its final development. It is, however, certain
that up to a late stage of growth, when the male Okapi is nearly
of full size, the ossicone has not penetrated the integument with
its tip, and that there is no indication of the polishing of the
ossicone’s tip nor of transverse fissures caused by absorbent
ingrowths of soft tissue. This is demonstrated by the ossicone of
a male* Okapi of nearly full growth (the last molar of the upper
series being not yet in use and the premolars only recently
having superseded their deciduous fore-runners), which is illus-
trated in Plate VII. This specimen belongs to a very perfect
skeleton obtained, together with the skin, by Major Powell
Cotton in the Ituri Forest, which is now in the British Museum
(Natural History) at Cromwell Road. The ossicone figured is
that of the right side. It is larger than that of the left side,
weighing 31:45 grammes as against 28°15 grammes scaled by the
left ossicusp. This asymmetry of the ossicusps of the Okapi, and
a difference in the direction of the slope of these structures when
right and left sides are compared, is to be observed in skulls of
adult male specimens, and was mentioned by me in my memoir
of 1901.
The ossicones in Major Powell Cotton’s specimen have not yet
* Major Powell Cotton ascertained that this specimen is a male, by an examination
of the genital organs, and the skin prepared under his direction retains the external
genitalia.
1907. | RUDIMENTARY ANTLERS IN THE OKAPI. 133
ankylosed with the frontal bone, but show an expanded base wath
radiating trabecular structure of the bony material (see fig. 2,
PV Uh: ys a peculiarity of surface which is repeated by the enlarged
area of frontal bone upon which they rest—a condition which
occurs also in young Giraffes. As shown by the drawings in
Plate VIIL., there is no evidence in this ossicone of polishing or
sharpening of the apex. The rough longitudinal grooves and
furrows are continued to that region, and there are no transverse
fissures. A vertical section (fig. 7, Plate VII.) of the apical region
shows a very dense bony structure, but no trace of ingrowths
from the surface. We may take it that this ossicone was still
entirely covered in by the vascular living integument, as is the
ossicone in the Giraffe throughout life. It furnishes us with
a stage immediately antecedent to the fixation of the ossicone by
ankylosis to the frontal bone, and antecedent to the breaking
through of the integument by the apex of the bony cone.
We may imagine the subsequent stages in the history which
connect this specimen with those figured in Plate VI. The rubbing,
polishing, and pointing of the ivory-like apex by use, the first
horizontal ingrowth of the lacerated investing integument, the
recession of that living investment after having established one
horizontal discontinuity or plane of disruption of the dense bone,
and the subsequent invasive ingrowth to form a second and by
repeated recession and ingrowth a third such plane of disruption,
and probably yet others.
In conclusion, I would poimt out that it is quite conceivable
that this cutting off of a series of antler-like tips from the
ossicone of the Okapi, is a process independently set up in this
Girafiid animal, having a similar physiological explanation to
that which applies to the similar process familiar to us in the
Cervide.
The different views which may be entertained, in the present
state of our knowledge of the facts of embryology and early growth,
as to the inter- relations of the ossicones of the Giraffidee, the
antlers of the Cervide, and the bony horn-cores of the Bovide,
are briefly stated in the preceding memoir on the “‘ Origin of the
Lateral Horns of the Giraffe,” and to this I would now refer the
reader.
[April 1907.—I add here text-figures (54 & 55, p. 134) of the
ossicone of the specimen of Okapi brought home by Capt. Boyd
Alexander from the Welle River and presented by him to the
National Collection. There are many interesting features about
the skin and skull which we owe to Capt. Boyd Alexander, and
these I hope to describe hereafter. The horn or ossicone is that
of an animal a very little younger than Major Powell Cotton’s
(as inferred from the dentition). It is intermediate in size
between that drawn in text-fig. 50 and that figured in Plate VII.
Its bony substance is much less dense and ivory-like than that of
134 ON RUDIMENTARY ANTLERS IN THE OKAPI. [ Feb. 5,
Text-fig. 54. Text-fig. 55.
Text-fig. 54.—Drawing of the right ossicone (not yet ankylosed) of the Okapi
brought by Capt. Boyd Alexander from the Welle River.
The relative age of this Okapi is indicated by the fact that whilst the deciduous
molars are still in place in every socket, yet they are very much worn and all
the true molars in both upper and lower jaw are “through ” and moderately
worn.
Text-fig. 55.—Section through the tip of the same specimen to show the incomplete
trabeculated ossification, contrasting with the dense ivory-like ossification of the
ossicone of Major Cotton’s specimen (Plate VII.).
The relative age of this Okapi is shown by the fact that the deciduous molars are all
shed and the premolars moderately worn, whilst the molars are all “ through ”
and moderately worn.
the larger and older ossicone, whilst less spongy than that of the
smaller and younger one. |
EXPLANATION OF THE PLATES.
Prater VI.
Figs. 1 to 8 represent various views of the “horn-tips” of the ossicones or
ossicusps of the Okapi in the Royal Scottish Museum, Edinburgh, of
the natural size (Specimen A).
Figs. 1, 4, 5, and 7 are views of the left horn-tip.
Figs. 2, 3, 6, and 8 are views of the right horn-tip.
Fig. 1. Left ossicone, outer side; y, depression due to shedding of a piece of the
bone.
Right ossicone, inner (mediad) side; x, cup-like cavity due to shedding of a
portion of the bone.
. Right ossicone, outer side.
. Left ossicone, inner (mediad) side; y as in fig. 1.
. Left ossicone, from behind and below.
. Right ossicone, from behind and below.
Left ossicone, from in front and above.
Right ossicone, from in front and above; «x asin fig. 2.
Figs. 9 to 12 represent two views of the “horn-tips” of the ossicones of a
specimen of Okapi lent by Messrs. Rowland Ward and now in
America (Specimen B).
Fig. 9. Left ossicone, outer side.
10. Right ossicone, outer side.
11. Left ossicone, from in front and above.
12. Right ossicone, from in front and above.
bo
WADE oo
1907. ] ON A NEW AMAZONIAN TREE-FROG. 135
Prate VII.
The figures illustrate the ossicone of the right side of Major Powell Cotton’s
male Okapi.
. The specimen of the natural size, seen from the right.
. The base of the ossicone, held by soft tissue to a similarly madrepore-like
surface of the frontal bone. Natural size.
3. Surface of the horn-tip, left side, enlarged to twice the natural size to show
the absence of polishing and of transverse fissures. The ossicone was com-
pletely covered by integument ; it had not been “ cut ” or emerged.
. Similar view of the right side.
. Similar view of the horn-tip from in front.
. Similar view from behind.
The horn-tip has been sawn through so as to remove the right-hand moiety
of the tip. The extremely dense, ivory-like character of the bone of this
region is thus demonstrated, and the absence of horizontal or other pene-
as fissures (compare and contrast with the text-figures, especially text-
g. 52).
doe
TO OE
4. Description of Hyla resinifictriv Goeldi, a new Amazonian
Tree-Frog peculiar for its Breeding-habits. By Prof.
Dr. Emit A. Gorip1, C.M.Z.8., Director of the Para
Museum.
[Received January 21, 1907. |
(Text-figures 56-59.)
In its warty skin this remarkably fine Tree-Frog resembles Hyla
tuberculosa Giinther, Hyla tawrina Steindachner, and Hyla venu-
losa Laur. It is most closely related to the last.
Length of a male, 8 cm. from snout to vent.
Head semicircular. Space between the two nostrils slightly
concave. Nasal region descending abruptly to the border of the
mouth, almost at right angles to the frontal plane. Canthus
rostralis running in a curve, rounded off. Nostrils, seen from
above, forming slight prominences. Choane large. YVomerine
teeth in two rows, forming an angle pointing forwards. Tongue
heart-shaped.
Tympanum very distinct, moderate sized, rather smaller than
the eye. Inthe male distinctly prominent vocal sacs between the
rictus of the mouth and the insertion of fore-legs.
Fingers III, IV, and V connected about half their length by
a web; no perceptible rudiment of thumb. Finger-disks large, a
little smaller than the tympanum. Outer border of arms without
folds of the skin. When the hind legs are stretched forward, the
tibio-tarsal articulation reaches the eye or the border of the mouth.
Heels without appendage.
Tubercles of sole of foot not so prominent as in Hyla taurina.
Colour greenish-yellow with blackish-brown markings: a brown
trapeze-shaped field between nose and anterior border of eyes; a
light-coloured, broad band running from one eye to the other,
the width of the eye, anterior bordering line straight, posterior
line slightly curved backwards; a large dark field covering all the
dorsal region, laterally rmmning down to the insertion of the fore
136 DR. E. A. GOELDI ON A [ Feb. Ds
legs, posteriorly leaving in the sacral region a triangular light-
coloured space. Between the insertion of the hind legs, situated
in the median line, there may be a small dark ring with a central
dark point (text-fig. 57 A). Dorsal area either entire or laterally
constricted about the middle (as in a fourth specimen not figured),
or divided in two isolated parts by a light-coloured cross-band
(text-figs. 56 and 57 B). The dark dorsal blotches are thickly
studded with thorn-like elevations (pointed warts) with a light-
coloured apex (text-fig. 56), more thickly agglomerated in the
Text-fig. 56.
Hyla resinifictrix, male (nat. size).
parotoid region (left side 13, right side 22, mm one individual,
and also in the posterior part 22+ 24 in one individual), besides
an abundance of smaller elevations and diminutive granulations.
A brown band around the upper arm, another around the fore-
arm, very well pronounced (with some lightish spots, imitating
the light central wart-spots of the back, but not raised); across
the hand several narrower bands. Legs: thigh with two bands;
lower leg with two broader bands, with the same whitish spots as
above mentioned ; across the ankle and foot several narrower bands.
Forehead with some dark, roundish spots.
Upper border of mouth dark-marbled.
1907. | NEW AMAZONIAN TREE-FROG. 137
Ventral surface light greenish-yellow, granulated and turning
warty about the chest. Upper-arm band on the anterior side
with two or three light spots; some light yellow elongated spots
at the anterior border of the vocal sacs. Border of lower lip
as well as the finger-disks of a delicate green.
From a side view with the extremities folded, on the anterior
extremity the dark bands on upper and lower arms appear to be
continuous with the posterior border of the anterior large dark
dorsal field; just as in the posterior extremity the dark bands
nearest to the knee (femoro-tibial articulation) coicide with the
posterior border of the second large dorsal field, while the bands
nearest the heel (tibio-tarsal) coincide with the round sacral spot
(text-fig. 57 A), the central spot forming with the lateral bands
and dots a fan-like arrangement (confer also text-fig. 57 B).
Text-fig. 57.
AX< B.
Hyla resinifictrix (diagrammatic).
Comparison of the four specimens available shows a considerable
extent of individual variation in the details of ornamentation.
However, it is easily seen that certain general features persist
fairly well, the most salient among them certainly being the
broad light transverse band on the head, running from one eye
to the other.
The iris is golden, with a horizontal and a vertical black bar,
forming a cross*.
This large and strikingly coloured Tree-Frog presents a most
curious novelty in its breeding-habits. Some years ago Mr,
Boulenger, of the British Museum, published in an interesting
* As described in H. venulosa by Boulenger, Ann. & Mag. N. H. ser. 5, vol. x.
p. 327 (1882).
138 DR. E. A. GOELDI ON A [ Feb. 5,
article a tabular synopsis showing the different incubation-habits
among the Batrachia. In this synopsis a unique position is held
by the tree-frog Hyla palmata, which, according to observations
made by me in the Serra dos Orgads (Rio de Janeiro), and
published in the ‘Proceedings’ of the Zoological Society, 1895,
forms breeding-bowls of the mud in shallow ponds.
Text-fig. 58.
Breeding-basin of Hyla resinifictrix, (A) side view and (B) section
(diagrammatic).
A still more peculiar eccentricity is presented by this beautiful
new Amazonian tree-frog, Hyla resinifictrix. Inhabiting the
virgin forest, it chooses certain tall trees for its dwelling, where 1t
takes possession of a hollow branch (text-fig. 58, A and B), and
constructs there as a nursery a good-sized basin of resinous sub-
stances, with a central depression (text-fig. 59). As is well known,
1907. | NEW AMAZONIAN TREE-FROG. 139
water and other liquids are preserved fresh in vessels varnished
with pitch, and in like manner the rain-water which fills these
resinous breeding-bowls presents excellent conditions for the
hatching and development of the eggs and tadpoles, such as shade
and freshness of water without contamination of decayed wood.
Without having had as yet the good fortune to verify it by direct
observation, I have abundant reason to suppose that the develop-
ment stage of the tadpoles is exceedingly brief, analogous with
the case of Hyla goeldii Boulenger in the Serra dos Orgads, Zyla
venulosa in Para, and others.
Text-fig. 59.
Breeding-basin of Hyla resinifictrix, seen from above (from a photograph).
One very interesting feature is the fact that this Amazonian
tree-frog goes in search of the material with which to build the
basin, and chooses for the purpose odorous resins which drop from
the bark of certain trees, such as the aromatic “ breo-branco ”
(Protium heptaphyllum) and others.
Although the resin of the “cunnuart” is well-known to the
Indians and mixed races in the Amazonian valley, constituting a
commodity much sought for and of high price, the tree-frog
itself was entirely unknown to all except the genuine forest-
dwelling Indians. In spite of strenuous efforts it took me more
than ten years to get on the track of this most mysterious
Batrachian, and if finally my efforts were crowned with success,
it was largely due to the friendly aid of the Tembé Indians at the
Mission of Santo Antonio do Prata, at the River Maracana (interior
of the State of Para), by the kind interest of Frei Daniel de
140 MR. OLDFIELD THOMAS ON MAMMALS | Feb. 5,
Samarate, Director of the Mission. With their help I succeeded
in obtaining several individuals, one of which ( 3) at least is still
alive after spending nearly two years in a terrarium properly
fitted up for it. Last year it gave me frequent opportunities to
hear its voice, which is surprisingly strong, and sounds as “ queng-
queng” three or four times repeated.
The local name ‘ cunnuart,” evidently onomatopeeic, is formed
by contraction of two Indian words ‘‘cunha=wife” and “art=
toad”; the Indians say that this tree-frog always calls for the
female when the moon shines.
5. The Duke of Bedford’s Zoological Exploration in Hastern
Asia.—III. On Mammals obtained by Mr. M. P.
Anderson in the Philippine Islands. By OLDFIELD
Tomas, F.R.S., F.Z.8.*
[Received February 5, 1907. |
In the early part of last year, after making the Korean
Collection described in a previous volume of our Proceedings *,
Mr. Malcolm Anderson paid a short visit to the Philippines, but
was unfortunately attacked by fever, and after a gallant attempt
to fulfil the object of his trip, was compelled to return to more
northern and healthier latitudes.
The chief object of Mr. Anderson’s visit to the Philippines was
to obtain series of the interesting mammals discovered in
Mindanao by Dr. E. A. Mearns, as the mountain fauna of this
island was only represented in our National Nuseum by the
duplicates from Dr. Mearns’s collection which the authorities of
the United States National Museum had been good enough to
send us. But these of course did not include any of the various
new genera and species which had been described by Dr. Mearns
on single specimens or on small series, and we therefore hoped
that Mr. Anderson might be able to obtain some of them for us.
Owing to Mr. Anderson’s illness the collection is quite small,
only consisting of 16 specimens belonging to 6 species, but one of
these proves to be new, while all are of interest as fillmg up gaps
in our series.
1. URoOGALE CYLINDRURA Mearns.
3. 756, 763. 92. 753, 761, 762. Mount Apo, Mindanao,
3000'—4000'.
Tam very doubtful if this form is at most more than a local
subspecies of U. everetti$, described from Zamboanga. The type
of the latter had been skinned after preservation in spirit, and such
slight colour differences as there are may be due to this cause.
* [The complete account of the new species described in this communication appears
here; but since the name and preliminary diagnosis were published in the
‘ Abstract,’ the species is distinguished by the name being underlined.—Ep1Tor. |
+ P.Z.S. 1906, p. 858.
{ See Mearns, P. U.S. Nat. Mus. xxviii. p. 425, 1905.
§ Tupaia everetti Thos. Ann. Mag. N. H. ser. 6, ix. p. 250, 1892.
1907.] FROM THE PHILIPPINE ISLANDS. 141
2. ScIURUS MINDANENSIS Steere.
3. 764. Mount Apo, Mindanao. 3000’.
3. Mus sp.
2. 748, 750. North Central Mindanao. 1100’, 3000.
@. 754. Mount Apo, Mindanao. 3000’.
A vat of the ratéus group. Dr. Mearns describes no less than
three species of this group, but does not seem to be aware that
the presence of spines in the fur in these rats 1s a most variable
character, depending in all on age, and in some on season, the
spines being shed and regrown periodically.
4, Mus spacosus Mearns.
Bullimus (g.n.) bagobus Mearns, P. U.S. N. M. xxviii. p. 450,
1905.
@. 755. Mount Apo, Mindanao. 3000’.
I fail to see any sufficient reason for the creation of a special
genus to contain this species. The small supplementary cusps on
the lower molars, on which Dr. Mearns mainly founds the genus,
are not only present in many Malayan species usually referred to
Mus, but they are even quite well-marked, though small, in his
own specimens of Mus albigularis Mearns, also from Mindanao.
None of the other characters mentioned by him appears to me of
generic importance.
Tt is unfortunate that Dr. Mearns had not had experience of
the difficulties of Murine dental characters before venturing to
describe genera of this group. Had he had such experience I am
sure he would not have described Bullimus, nor would he have
based another genus (Limnomys) on a single specimen with teeth
“too worn to furnish characters distinguishing them from J/us,”
unless the other characters were of a far more striking nature
than appears from his account.
Mr. Anderson’s specimen of JZus bagobus, being a female,
enables me to state that the mamme number 1—3=8, a formula
only occurring to my knowledge in two other Hastern species
of Mus.
5. Mus vuntcant Mearns.
@. 752. North Central Mindanao. 4000’.
A member of the common and widely spread concolor-ephippium
group.
6. CRUNOMYS MELANIUS Thos.
Aosine, 12, At WOOT. 1. D (Wdelos 124)
3g. 751. North Central Mindanao. 3000’. 29 February,
1906. B.M. No. 7.2.2.14. Type.
A species with the general appearance of a blackish Akodon of
142 ON MAMMALS FROM THE PHILIPPINE ISLANDS. [ Feb. 5,
the A. caliginosus group, or perhaps more of a member of the
subgenus Melanomys.
Size rather larger than in C’. fallaaw of Luzon. Fur close and
fine, but rather stiff owing to the admixture of a number of fine
flattened spines; hairs of back about 6-7 mm. in length. General
colour above blackish brown, nearest to bistre of Ridgway
but darker; the under surface nearly ss dark as the upper,
without any line of demarcation, Individually the hairs of the
back are dull slaty basally, with buffy tips ; the spines slaty with
black tips. Ears very short, closely haired, brown. Whole of
limbs, including hands and feet, blackish brown. Tail short, very
finely scaled (scales about 18 to the centimetre), finely haired, the
hairs about 14 scales long ; uniformly blackish throughout.
Skull less flattened than that of C. fallax, but this may be due
to youth. The brain-case long, comparatively parallel-sided, in
continuation of the broad interorbital region. Supraorbital edges
with scarcely a trace of beading. Outer plate of zygoma-root cut
back just as in C. fallaw. Palatal foramina short, narrow in front,
broadened behind, not parallel-sided as in fallax.
Molars rather like those of a very simple type of J/ws, without
any supernumerary lateral cusps. M, and M, each with a small
median posterior cusp as in C. fallax.
Dimensions of the type, which is barely adult :—
Head and body 98 mm.; tail 68; hind foot (s.u.) 25; ear 13.
Skull—greatest length 28 mm.; basilar length 22 ; zygomatic
breadth 14; length of nasals 9°2; interorbital breadth 5-6;
breadth of brain-case 12°3; palatilar length 11; diastema 7;
palatal foramina 3°8 x 2:2; length of upper molar series 4:1.
Hab. and Type as above.
The characters of this remarkable species are a great puzzle,
and only add to the difficulty of assigning a proper position to
the type of the genus, Crunomys fallax. The single specimen of
the latter is very old, and in the worn-down state of the teeth it
was not clear whether they were or were not of hydromyine
structure. The teeth of the present animal are certainly not
typically hydromyine, but rather murine, while at the same time
it is possible that in wearing down they might acquire the slight
resemblance to hydromyine teeth shown by C. fallax. This being
the case, I have decided to describe the species, though with doubt,
as a Orunomys, but until younger specimens of C. fallax, or older
ones of the present form, are available for examination, it would
be advisable not to express any definite opinions as to the
systematic position of either. As a species C. melanius is at once
distinguishable by its blackish colour and heavier feet.
7. SUS sp.
4 skulls. Mount Apo, Mindanao,
P.Z.S. 1907. Pl. VIII.
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AGRIOTYPES OF THE Sf&RIPED TABEY.
1907.] ON ENGLISH DOMESTIC CATS. 143
February 19, 1907.
Sir Eymunp G. Lover, Bt., Vice-President, in the Chair.
The Secretary read the following report on the additions that
had been made to the Society’s Menagerie in January 1907 :—
The registered additions to the Society’s Menagerie during the
month of January were 108 in number. Of these 55 were
acquired by presentation and 11 by purchase, 41 were received
on deposit, and 1 was born in the Gardens. The total number
of departures during the same period, by death and removals,
was 163.
Amongst the additions special attention may be directed to :—
An Agile Gibbon (//ylobates agilis) from Borneo, purchased on
Jan. 17th.
A Wild Cat (Felis catus) from Inverness-shire, presented by
Mr. G. W. Henderson, F.Z.5., on Jan. 26th.
Two Superb Tanagers (Calliste fastwosa), a Crowned Tanager
(Tachyphowus coronatus), and two Scarlet Tanagers (Rhamphocelus
brasilius), from Brazil, purchased on Jan. 21st.
Dr. C. I. Forsyth Major, F.Z.8., exhibited remains of a Bear
from the superficial deposits of a cavern in the mountains of
Corsica, where Bears, though now extinct, were formerly nume-
rous, at least up to the sixteenth century. Despite the fact that
no truly fossil Bears were as yet known from Corsica, Dr. Forsyth
Major considered the Corsican Bear to have been autochthonous,
whilst in his opinion the recent Mammals of Corsica (and Sardinia)
had been, almost without exception, introduced by human agency.
In any case they could not be adduced as proofs of a recent
connection of those islands with either of the nei ghbouring
continents.
The following papers were read ;—
1. On English Domestic Cats. By R. I. Pococn, F.L.S.,
F.Z.8., Superintendent of the Zoological Society’s
Gardens.
[Received February 5, 1907.)
(Plates VIIIT_—-X.* and Text-figure 60.)
1. The Classification of English Domestic Cats.
Domestic Cats in Great Britain are classified by the National
Cat Club under two headings :—(1) Short-haired ; (2) Long-
haired, otherwise called “Persians” or ‘“ Angoras,” The
“ Persians” are subdivided according to colour into “ Blacks,”
* Foy explanation of the Plates, see pp. 167, 168.
144 MR. R. I. POCOCK ON [ Feb. 19,
“¢ Smokes,” “Sables,” “‘ Blues,” “‘ Oranges,” “‘ Creams,” “¢ Whites,”
“Tabbies,” ‘“ Tortoise-shells,” &¢. The ‘Short-haired” are
similarly distinguished, after the elimination of certain types like
“Siamese,” “ Manx,” and “ Abyssinians,” to which special classes
are assigned *.
From the exhibitors’ and breeders’ points of view this arrange-
ment has much to recommend it, and probably supplies the most
feasible and satisfactory method of classifying the various breeds
that are set side by side for comparison. The animals must be
sorted out upon some basis, and it would perhaps be impossible to
suggest a substitute that would meet the requirements of fanciers
equally well.
But, from the standpoint of affinity and descent, I think it is
open to the criticism that a primary impor fance is given to
characters like the length and thickness of the fur, the tint of the
ground-colour, and the absence of the tail, which can be shown to
have no great systematic value; whereas a quite subsidiary signi-
ficance is attached to the nature of the pattern of the stripes, a
character which should be of the greatest moment in differen-
tiating the breeds, if confidence be paced i in the analogy supplied
by existing species of the genus Felis.
It may fairly be asked, however, why a greater taxonomic value
is claimed for the “pattern” than for the three characteristics
mentioned above upon which fanciers establish their breeds. The
answers are briefly these :—
1. The shortness of the hair in tropical Leopards and Tigers as
compared with those that come from colder countries shows that
no great systematic importance can be attached to length of
coat. The difference between the extremes of long- and short-
haired Domestic Cats is admittedly much greater than that between
long- and short-haired Leopards or Tigers: but all gradations
between ‘“ Persian” and ordinary Cats exist, so that no hard-and-
fast line can be drawn between them; and probably no one
doubts that the luxuriant growth characteristic of the former
breed has been preserved and increased by artificial selection.
In the matter of coat, the adaptability of Domestic Cats to
changed conditions is proved by the evolution of a thick-haired
breed in the Pittsburgh refrigerators 7 and in the arctic island of
St. Paul t, and by the alleged shortness and stiffness of the fur in
Domestic Cats from Mombasa in Hast Africa §.
2. The conclusion that the shortness or complete absence of the
tail in so-called “ Manx Cats” has been brought about by selective
breeding from favoured sports must be regarded as beyond
dispute. There is no such thing as a tailless species of Felis
amongst fossil or recent forms. The tail is short in Lynxes, but
* ©Cats and all about them,’ 1902; and ‘The Book of the Cat,’ 1903, -by
Frances Simpson.
t Quoted from Lydekker, ‘Cats, &c.’ p. 159 (1896) (Allen’s Nat. Library).
* See H. C. Marsh, ‘Darwinism,’ p. 21 (1883).
§ Darwin, ‘ Animals and Plants under Domestication,’ p. 58 (ed. 1905).
il 907.| ENGLISH DOMESTIC CATS. 145
it cannot be claimed that an intermixture with any known
species of Lynx has contributed to the abbreviation of the tail in
the “ Manx.” When and where this breed arose has been much
debated. Some have suggested Spain as its original country,
others China. But in all probability the “ sport” has appeared
independently and been preserved by selective crossing on many
occasions and in many places.
The liability of the tail to modification in Domestic Cats is
shown by the frequency with which it is kinked or twisted as
well as shortened in Bur mese, Siamese, and Malaccan specimens ;
and by the stunting it acquired in a few generations in the Cats,
above alluded to, that were bred in the frigid climate of St. Paul’s
Island and in the cold-storage warehouses of Pittsburgh.
3. With respect to tint or the ground-colour as a whole, irre-
spective of pattern, it must be remembered that albinotic and
melanotic “sports” may arise in almost any group of Mammals.
Amongst the Cats, albinos appear to be rare in a state of nature,
probably in part on account of a correlated inherent delicacy 1 in
organisation, accompanied possibly by defective sight or hearing ;
probably in part on account of the conspicuousness of the
coloration making capture of prey and escape from enemies,
especially during cubhood, difficult. There are one or two
records, however, of albino Tigers—the species, be it noted, in
which the young after leaving the mother’s protection are more
capable of taking care of themselves and less liable to attack than
any other Cat, with the possible exception of Lions. Melanisms
are far more frequently met with in the genus. Black Leopards
are familiar to all; black Jaguars are not uncommon, and black
Tigers, Caffre Cats, and Servals have been recorded,
In other species the tint may be dimorphic, dark grey or dark
brown and red or chestnut examples being about equally abundant,
as in the 8S. American Felis jaguarondi, the West African Velis
aurata, the Oriental F’. temmincki, and the Bornean /. badia.
Thus in the genus Felis colour-variation in a state of nature
may depart from the normal in the direction of “black,” or
“ved,” or “ white,’ without respect to locality. Moreover,
geographical variation also attesting inherent instability of tint
is met with. The greyness of the Persian Leopard as compared
with the rich tawny yellow of South Indian specimens is a case
in point.
The tints of Domestic Cats themselves establish the same con-
clusion. Between “ blacks” and “ whites” all intermediates seem
to exist. ‘ Blues” are self-coloured slate or lavender-grey cats
and appear to be merely stages in the direction of “blacks.”
“Smokes ” are darkish grey or blackish cats with the basal part
of the hair white. ‘“‘ Silvers” are the “‘ palest conceivable edition ”
of Smokes. ‘ Reds,” “ Yellows,” and ‘‘Creams” are variations
from the normal in the direction of ‘‘ Whites.”
4. On the other hand, notwithstanding individual and local
variations, the pattern formed by spots or stripes in existing
Proc. Zoou. Soc.—1907, No. X. 10
146 MR. R. I. POCOCK ON [Feb. 19,
species of Felis is on the whole constant. Stripes may break up
into spots or spots may run together to form stripes, or both spots
and stripes may be altogether fugitive. Yet even in extreme
and rare cases of this nature, the general “character” of the
pattern, when detectable, remains the same, and there is abund-
ance of evidence that the animals breed true to the local type.
Nor, so far as I am aware, is there any reason to suppose that
dimorphism in pattern ever occurs or has ever occurred im any
species of the genus Felis.
It is needless to say more in support of the contention that if a
decided difference in the “ patterns” of Domestic Cats exists, it
must be regarded as furnishing a surer basis for their classification
than the length of the hair, the tint of the coat, or the stunting
of the tail.
It may also be claimed with assurance that the pattern supplies
a more important clue to the ancestry of Domestic Cats than the
features just mentioned.
Probably no one will dispute that all breeds of Domestic Cats
have been derived from one or more than one ancestral type that
was marked with bands or spots. This opinion is supported by
two considerations. The first is this: spotting or banding of one
type or another characterises the great majority of the existing
species of Cats, using this term in its broad sense as co-extensive
with the genus Felis*. A few self-coloured Cats, like Lions,
Pumas, Caracals, and others, exist ; but their descent from striped
or spotted forms is attested in most cases by the presence on the
cubs + of markings which are subsequently lost or by indications
of them, especially on the legs or lower parts of the body, in the
adults.
The second pertinent point is the prevalence of stripes forming
a pattern of one kind or another, both in “ Persians,” “ Short-
haired,” and “ Manx” breeds, and the difficulty breeders experience
in eradicating these markings in their efforts to preserve a
particular self-coloured type. Frequently at all events the so-
called “ blotched ” pattern can be detected in certain lights even
in “ Whites ” and “ Blacks,” the two varieties which stand at the
extremes of the colour-mutations of the diverse domestic breeds.
Assuming, then, that domestic breeds are descended from one or
more than one striped or spotted species, we may safely set aside
the self-coloured forms as derivatives and consider only the
striped or spotted types in looking for the origin and for a reliable
basis for the classification of Domestic Cats.
Domestic Cats in which the markings on the skin form definite
patterns are called comprehensively “ Tabbies.” Of “Tabby”
Cats, as fanciers well know, there are two kinds, the “striped ” and
the “blotched.” These are not, however, regarded as different
* TI purposely ignore here the subgeneric divisions of Felis adopted in Trouessart’s
‘Catalogue of Mammalia,’ because I cannot admit that these connote, at least in all
cases, natural assemblages of species.
+ Young Caracals are not spotted or striped, but resemble their parents.
1907. | ENGLISH DOMESTIC CATS. 147
breeds. Nevertheless the patterns appear to be fundamentally
distinct from each other in the sense that the differences between
them are not differences of degree, but of kind. Without
assuming the existence in the past of a number of intermediate
stages which do not appear to exist in the present, it 1s impossible
to reduce them to a common plan and it is difficult to see how
one can have arisen from the other unless per saltwm*.
One or the other of these patterns, when the pattern is traceable
at all, may be seen in Cats of the short-haired, long-haired, or
Manx breeds, whether the ground-colour be grey or red, or black
or white, or any other tint. Neither at Cats’ Homes nor at Cat
Shows nor in the streets have I seen one cat out of the hundreds
observed in which the pattern could not be assigned at once to
one or the other of these types. Here, then, is evidence for the
existence, side by side, of two fundamentally distinct kinds of
“‘ Domestic Cats.” These I propose to regard as species, trusting
to the analogy supplied by wild forms of the genus felis and
knowing no reason for thinking, much less proof of the fact, that
one has been derived from the other, either as a sudden sport or
by gradual modifications under the influence of selective breeding
or by inter-breeding with any wild species of the genus.
The two types are described in some detail below (see pp. 151-
153). They may be very briefly diagnosed as follows :—
a. Sides of the body, from the shoulder to the root of the tail,
marked with narrow wavy vertical stripes which show
a tendency, especially on the thighs, to break up into
spots ; no broad latero-dorsal stripe ...... Striped “ Tabby.”
b. Sides of the body marked with three usually obliquely
longitudinal stripes forming the so-called “ spiral,”
“horseshoe,” or ‘ circular” pattern of fanciers; a
broad latero-dorsal stripe on each side of the narrow
median spinal stripe ............ sdsroialalasic'os Blotched “‘ Tabby.”
It is difficult to ascertain from the writings of earlier natura-
lists whether they were familiar with these two types of Cats or
not. There is certain evidence that the blotched or true “Tabby”
Cat was domesticated in Sweden and known to Linnezus as early
as 1746. And that the Striped Cat was also domesticated in
Kurope at a still earlier date is proved, I think, beyond doubt by
the figures published in Gesner’s Hist. Anim. p. 345 (1551), and in
Johnston’s ‘Quadrupeds,’ pl. Ixxil. p. 126 (1657). If these and
later post-Linnean authors distinguished the two Cats, it must be
inferred that they attached no significance to the difference of
pattern, but regarded it as of the same value as the difference of
colour and as the asymmetrical blotching of piebald specimens.
Pennant, for example, speaking of the Tame Cat, says that it
* The differences between these two types are described and figured by Mr. T.
S. Rope (‘ Zoologist,’ 1881, pp. 353-357). Perhaps the omission of this paper from
the ‘ Zoological Record’ may explain in part the want of consideration the facts
have received from most recent zoological writers on this subject. .
10
148 MR. R. I. POCOCK ON [Feb. 19,
differs only ‘‘in color and some other trifling accidents” from the
Wild Cat (British Zoology, vol.i. p. 94, 1812), and Kerr describes
the stripes on the sides of the Wild Cat as being “perpendicular
or spiral” (Anim. Kingd. p. 152, 1792), thinking apparently that
the blotched ‘‘ Tabby” was a domesticated form of the European
Wild Cat and that the shape and direction of the stripes were
subject to variation and of no particular moment.
Even the writings of later authors leave them open to suspicion
on this point. Gray, for example, pointed out that a Domestic Cat
brought by Darwin from 8. America was remarkable for its striking
likeness to the Caffre Cat. But there is nothing surprising
in this if, as I suppose from the description, the Cat in question
was merely a slightly aberrant example of the Striped “ Tabby”
breed. This Gray could hardly have failed to detect had he
been familiar with the Domestic Cats of London. Again, since
Blanford and Mivart both regarded Indian specimens of the
“Striped breed,” known as /’. torquata, as examples of a genuine
wild species, it may be inferred that they were both ignorant of
the fact that Cats inseparable from that type, as figured by Cuvier,
might be seen any day in London, where and at the time when
their volumes, below (p. 159, footnote) cited, were being written.
Setting aside breeders and owners of “ Fancy Cats,” who could
hardly be expected to appreciate the significance of the differences
of pattern above described, although well aware of their existence,
two scientific writers on Cats must be mentioned as clearly
apprehending the fact. One of these is Rope, whose paper,
published in 1881, has been already mentioned; the other is
that astute observer Blyth, who so long ago as 1845* pointed
out that the two types of pattern are found in the Domestic Cats
of Europe. In an additional note on the subject published
in 18567, he spoke of the true Tabby pattern as being possibly
a “ modification (and a very remarkable one) of the markings of
the wild F. sylvestris of Europe.”
Rope apparently did not know of Blyth’s papers. All the
more remarkable is it therefore that both authors describe this
Tabby as being marked with pale streaks on a dark ground; and
this description was repeated by Hamilton in his notice of Rope’s
paper. It is obvious, however, that no Cat can be scientifically
described as marked in this way. Hence it is possible that the mis-
leading terminology employed by Blyth, Rope, and Hamilton may
have suggested to the readers of their writings that the blotched
“Tabby” markings belong to the same category as the piebald
or skewbald, usually asymmetrical coloration of black-and-white
or brown-and-white cats, dogs, horses, cattle, and other tamed or
domestic mammals infected with either melanism, albinism, or
erythrism, or with any two of these taints, or with the three
combined. This may partly account for the fact that the difficulty
of deriving the blotched Tabby pattern from that of /. ocreata or
* Journ. As. Soc. Bengal, xiv. pt. 1. pp. 342-343.
“+ Op. cit. xxv. p. 443.
Urey ENGLISH DOMESTIC CATS. 149
F. sylvestris, and of accounting for the origin of the breed, seems
never to have been fully realised, or, if realised, never seriously
faced. And doubtless a contributory cause to this result has
been the artificial character of the classification of Domestic Cats
adopted by fanciers, which gives emphasis to valueless features
and obscures the fundamental importance of the pattern.
2. On the Characters and Names of the Two Types of Domestic Cat
and the Name of the European Wild Cat.
In the foregoing and following pages the European Wild Cat
is referred to as Felis sylvestris. This requires explanation, since
the species in question has, by almost common consent, been
hitherto called Felis catus.
In my opinion there is no possibility of evading the conclusion
that the Cat to which Linnzus gave the name catus was not the
European Wild Cat, but the Domestic Cat of the blotched Tabby
kind. In the 10th edition of the ‘Systema,’ 1758, accepted as the
starting-point in zoological nomenclature, Felis catwus is character-
ised in such terms as to leave no room for doubt on this head. In
the first place, there is a back reference to the ‘ Fauna Suecica,’
1746, where of Felis catus it is said “habitat in domibus; cicurata:
exotice originis.” In the second place, the diagnosis in the
‘Systema’ runs as follows :—‘“ Felis caudata, elongata ; corpore
fasctis nigricantibus ; dorsalibus longitudinalibus tribus ; laterali-
bus spiralibus.” 'The spiral lateral stripes are obviously those that
form the so-called “ circle” or “horseshoe” characteristic of the
blotched ‘‘ Tabby ” ; and the three dorsal longitudinal stripes are
also typical of that animal. This description is totally inapplicable
to the European Wild Cat, which moreover does not occur in
Scandinavia, and was apparently unknown to Linneus, except
from books. It is equally inapplicable to the “Striped” race of
Domestic Cats. On the other hand, it exactly fits the blotched
Tabby Domestic Cat, and was quite likely taken from a specimen
lying on Linneus’s hearthrug at the time.
There is no doubt from the context and the bibliography in
the 10th ed. of the ‘Systema’ that Linnzus believed this Cat
and the European Wild Cat to be the same species of animal ;
but that error does not in any way affect the certainty of the
conclusion that he knew enly the domestic blotched “Tabby” and
described it as catus. ;
The name “ domestica” must, I think, be discarded as a
synonym of caétus, since both Schreber and Erxleben, the two
earliest post-Linnean authors to use that term, quote under the
heading domestica Linneus’s diagnosis of catws. And since all
the so-called Tortoise-shell Cats that I have seen belong to this
type, the name hispanica Erxl. may also be placed amongst the
synonyms of catus ; vulgaris Fischer is also a synonym.
The name ‘catus,” then, is no longer admissible for the
European Wild Cat. For this species there are two names
150 MR. R. I. POCOCK ON [ Feb. 19,
available, which were apparently published in the same year ;
and between these a choice has to be made. These are :—
ferus Erxleben, Syst. Regni Anim. i. pp. 518 & 522 (1777).
sylvestris Schreber, Siug. ili. p. 397 (1777), pl. evii. a. (date 2)*.
The synonymy of felis catus ferus published by Erxleben
shows that he followed Linnzus in regarding the European Wild
Cat as what may be called the “agriotype” of the various
domestic breeds, such as domestica, angorensis, hispanica, and
cerulea, described on pp. 520-522; and the excellent diagnosis
of the Wild Cat published on p. 522 beginning Catus Yerus major,
proves that he was acquainted with that species, and applied the
name ferus to it, and not to feral examples of a domestic breed.
But for the following reasons I think sylvestris should be
preferred :—(1) Schreber adopted this name from Brisson’s Regn.
Anim., Quadr. p. 265, which, being published in 1756, is pre-
Linnean so far as nomenclature is concerned. Nevertheless, the
diagnosis of /, sylvestris suggests that Brisson distinguished this
Cat from the domestic catus of Linneus. (2) Erxleben quotes
both Brisson’s and Schreber’s works in his _ bibliographical
synonymy of /. catus ferus. Of Schreber’s work he correctly
cites pl. evii. A. inscribed Felis Catus Linn. ferus. Hence I infer
that he saw or knew by hearsay of this plate. I cannot, however,
find proof that it was published, properly speaking, before the
issue of the text in 1777. Since, therefore, the name sylvestris
in the text has page priority over the name /erws on the plate,
preference should be given to the former and the latter regarded
asasynonym. (3) The name sylvestris has been used for this
Cat by some modern writers, notably by Blyth and Hamilton.
For these reasons I think it advisable to adopt sylvestris instead
of ferus as the specific name for the European Wild Cat.
Apart from the convenience of having a name for a Cat which
not uncommonly occurs feral in the tropics, the determination of
the correct name for the “Striped” Domestic Cat is a matter
of no great moment. Moreover, it is practically impossible to
settle with certainty which name should be chosen out of the
many given by earlier writers to various domestic breeds in
different parts of the worldt. In addition to domestica and
hispanica, already referred to, the following are the most im-
portant :—ceruwea Erxl. for the ‘“ Blue Cat,” which seems to be
a pale or incompletely melanistic sport perhaps of the striped,
perhaps of the blotched type; angorensis Gmel., for the long-
haired breeds, which were no doubt originally of the “ Striped ”
type; ruber id., for a red variety with a dark dorsal stripe ;
sinensis Kerr, for a Chinese race alleged to have pendulous ears ;
* Kor discussion of the dates relating to Schreber’s work see Mr. C. D. Sherborn’s
paper, P. Z.S. 1891, pp. 588-589.
+ These names have to be catalogued to prevent their inadmissible use in a
different sense by later writers (see Sherborn, Index Anim. p. 187, 1902).
1907. ] ENGLISH DOMESTIC CATS. 151
aureus id, (=longiceps Bechst.), for a yellowish, short-legged, long-
headed, sharp-nosed breed, said to inhabit New Spain ; mada-
gascarensis id., for a Madagascar Cat with a twisted tail; striata
Bechst., for a black-striped “ Cyprian” Cat, which is diagnosed
as follows :—‘ Mit schwarzen Streifen auf hellem Grunde, die
auf dem Riicken gerade, auf den Schenkeln aber gekrimmt
sind” (Bechstein, Pennant’s ‘ Uebersicht, etc.’ ii. p. 679, 1800).
In a general way this description applies to both types of Cats
under discussion, and perhaps on the whole more clusely to the
“blotched ” than to the “striped” breed. But if the “Cyprus ”
Cat came originally from Cyprus, a conclusion by no means
justified by the epithet, it belonged in all probability, as did
the Angora Cat, to the “Striped” and not to the “ Blotched”
type*.
But whatever opinion may be held with regard to angorensis
and striata, there is no doubt that examples of this Cat, whether
feral or not, furnished types for the following forms :—
F. torquata F. Cuvier, Hist. Nat. Mamm. pl. 126 (1826),
recorded from Nepaul, Bengal, &c. (see infra, p. 165).
F. inconspicua Gray, Charlesworth’s Mag. N. H. p. 577
(1836).
Vip ee J. A.S. B. xv. p. 169 (1846), xvii. p. 247,
xxii. p. 581, from Candahar.
F. ocreata agria Bate, P. Z.8. 1906, p. 317, from Khania in
Crete.
The figure of F. torquata and the description of 2. huttoni
leave no room for doubting the identity of the Cats; and the
types of inconspicua and ocreata agria are in the British Museum
and have been compared by me with English examples of Striped
Cats.
This type (torquata) may be described as follows :—Ground-
‘colour typically iron-grey or yellow-grey. The four cephalic and
cervical stripes sometimes distinct, sometimes indistinct. The ad-
median cervical stripes not abbreviated on the anterior area of the
nape; the external cervical stripes not diverging from the middle
line on its posterior area. Dorsal area of body from the shoulders
to the root of the tail darker than the sides, the pigment often
resolvable into three narrow, almost contiguous stripes, two
laterals more or less interrupted, and a more complete median
which is usually continued down the middle line of the tail. On
the summit of the shoulder the lateral stripes are frequently
thicker and more heavily pigmented than they are posteriorly.
From the dark dorsal area on to the belly pass a number of
vertical wavy transverse stripes, which are usually more distinct
on the thoracic than on the abdominal region, where, as also on
the thighs, they are more or less broken up into spots, which may
* See Temminck, Mon. Mamm. i. p. 128 (1827); and Fischer, Syn. Mamm. p. 208
(1829).
152 MR. R. I. POCOCK ON [ Feb. 19,
or may not show signs of transverse lear arrangement. The neck
and the shoulders may be striped or merely speckled. The fore
and hind limbs are transversely barred; the former are typically
black behind up to the wrist and the latter up to the hock. The
tail also is transversely barred above, sometimes throughout its
length, sometimes only in its distal portion, the tip being black.
In specimens that I regard as typical of this Cat the pattern is
very similar to that of the European Wild Cat (7. sylvestris).
In others the transverse stripes break up into more or fewer,
larger or smaller spots. In others the spots are evanescent, the
fur being merely “ticked,” a gradation being traceable from the
first or sylvestris-type to the last, which are, I believe, the so-
called “ Abyssinian” and “Ticked” breeds of the fanciers. I
have obtained all these types from the Cats’ Home in Camden
Town, London, in the space of a few weeks, as well as a fine red
variety of the spotted kind. With the exception of the sylvestris-.
type, all these phases can be matched approximately in the series
of skins of /’. ocreata preserved in the British Museum.
Specimens which are intermediate between the ‘ spotted” and
the “ ticked” types—those, that is to say, in which the spots are
very small and closely set, giving a blackish appearance to the
skin except on the legs and ventral surface where the stripes are
apparent—deviate from the normal in exactly the same way as
the two South-African Leopards described by Dr. Giinther* differ
from normally spotted Leopards.
In the British Museum Collection there are skins of this
Domestic Cat from such widely separated localities as Crete,
India, Madagascar, Celebes, and Mexico. The specimen from
Celebes was shot on Bonthain Peak. It is noticeable that its.
stripes and spots are browner than in typical Kuropean examples ;
and in the skull the posterior portion of the nasals is more com-
pressed und the interparietal crest is higher and traceable as a
ridge all along the parietals. The narrowness of the nasals
recalls that of the skulls of two Siamese Cats that I have seen
(cf. infra, p. 163), and the height of the interparietal crest is
paralleled in the skull of a rufescent Indian Cat + which formerly
belonged to Mr. H. C. Brooke and was, I understand, a well-
known prize-winner at Cat Shows some few years ago. For
further information about torquata, see below, pp. 164-165.
The following description, taken from a number of skins, will
perhaps convey a tolerably clear conception of the plan (¢f. text-
fig. 60, p. 154) to which the patterns of F. catus, however
diversified in detail, are reducible :—
Ground-colour typically brownish or grey, frequently washed or
clouded with blackish. Head and face normally striped. ‘The
internal or admedian cervical] stripes abbreviated on the fore part.
* P.Z.S. 1885, p. 243, pl. xvi.; id. op. cit. 1886, p. 203, fig. See also W. L.
Sclater, Mamm. S. Afr. i. p. 36, fig. 10 (1900).
+ This Cat, Mr. Brooke tells me, was brought as a kitten from a hotel in Bombay.
[Oy a ENGLISH DOMESTIC CATS. 153
of the nape, and entering the anterior apex of a diamond-shaped
space (c.sp.), which is bordered laterally by the posterior ends of the
external cervical stripes and the anterior ends of a pair of broad
suprascapular stripes (szpr.sc.). The latter lie near the middle line
on the summit of the shoulder, and diverge forwards to meet at
an obtuse angle the posterior ends of the backwardly-diverging
external cervical stripes. From this angle stripes pass obliquely
forwards and downwards on to the sides of the neck and throat,
the posterior of them lying in front of the shoulder and forming
a throat collar. From the suprascapular stripe one broad vertical
stripe, the scapular stripe (sc.), passes over the shoulder and loses
itself in the uppermost of the transverse brachial stripes of the
fore leg. This scapular stripe forms the anterior border of the
postscapular space (p.sc.sp.), which is bounded behind by a second
broad vertical stripe, the postscapular stripe (p.sc.), descending
from the posterior end of the suprascapular stripe. Extending
along the middle of the spine from the posterior apex of the
cervical space there is a very narrow spinal or median dorsal stripe
(sp.), which passes down the middle line of the tail. Running
forwards on each side of this from the root of the tail to the
posterior end of the suprascapular stripe, which it frequently joins,
there is a very broad latero-dorsal stripe (/at.dors.). Beneath or
externally to this on the body there are three broad stripes
whose direction is obliquely longitudinal. The upper of these, the
supero-lateral (swp.lat.), forms a bold downward curve posteriorly,
while the inferior, the infero-lateral (inflat.), curves upwards
anteriorly. They thus partially circumscribe an elliptical or
subcircular area, in which lies the short and broad medio-lateral
stripe (med.lat.), which frequently has the form of a large spot
or blotch. These three stripes form the so-called “ horseshoe,”
“ circular,” or “spiral” mark characteristic of this type of
Domestic Cat. On the thighs there are broad transverse stripes,
the upper of which, the femoral stripe, extends obliquely down-
wards and forwards from a point beneath the posterior end of the
latero-dorsal stripe to a point beneath the downcurved posterior
end of the supero-lateral stripe. In the space between the three
stripes just mentioned lies a large spot or stripe, which frequently
fuses with the femoral stripe. On both fore and hind legs the
stripes thin out and die away towards the paws, which are typi-
cally black behind up to the wrist and hock. The tail is black at
the tip, and marked throughout with transverse black bars, which
are very broad where they touch the median caudal stripe.
The pattern of this Cat varies considerably in detail with
respect to the width and degree of fusion of the stripes. The
cervical and postscapular spaces are sometimes hardly apparent,
and the stripes may widen and fuse to such an extent that an
almost totally black Cat results. It is possible that the blackness
of some Cats is attributable to this process; but usually the
blackness is due to the melanism of the ground-colour, the
stripes retaining their normal width and being detectable under
154 MR. R. I. POCOCK ON [Feb. 19,
reflected light by the greater glossiness of the hair, The stripes
in white Cats are often visible in the same way. Tortoise-shell
variations of this Cat are nothing but partially erythristic or
erythro-melanistic sports, frequently with pronounced albinism
Text-fig. 60.
_MED. LAT, ------------
SUP. LAT. -------------=
LAT. DORS.
Diagram of the Pattern of the Blotched Tabby (Felis catus).
¢.sp., cervical space; p.se.sp., postscapular space; se., scapular stripe; swpr.sc., supra-
scapular stripe; p.sc., postscapular stripe; sp., spinal stripe; lat.dors., latero-
dorsal stripe; swp.lat., supero-lateral stripe; med.lat., medio-lateral stripe ;
inf.lat., infero-lateral stripe.
affecting particularly the legs, inferior areas, and even other parts
of the body. In simple cases the blotching may be seen to be
due to the red tint occurring only on portions of the stripes,
the remaining portions of which retain their normal black hue.
1907.] ENGLISH DOMESTIC CATS. 155
The result is a mixture composed of red and black stripes on a
yellowish ground-colour. But in more complicated cases the red
and black also invade the ground-colour in patches, either the
red or the black, but more commonly the black, predominating.
On these skins the original pattern is almost entirely obscured.
It is a well-known but none the less singular fact that Tortoise-
shell Cats are generally females and “ red” Cats generally males.
This circumstance is the foundation for the saying that the ‘red ”
Cat is the male of the “ Tortoise-shell ” ; but since the Tortoise-
shell is as much melanistic as erythristic, male black cats have
equal claims to be so regarded.
3. On the Characters of the European and African Wild Cats
(F. sylvestris and F. ocreata) and the Origin of the Domestic
Types (torquata and catus).
Criticism of the opinions of authors on the subject of the origin
of “‘ Domestic Cats” must be prefaced by the remark that, with
one or two exceptions, they failed to realise that an explanation
was required of the origin of two distinct kinds of Cats differing
so much from each other that no one would hesitate to regard
them as representing widely divergent species if they occurred in
a state of nature.
A great deal has been written on this topic. Opinions at one
time were nearly equally divided on the point, some authors
regarding the European Wild Cat (/. sylvestris), others the
Egyptian Cat (/’. ocreata) as the ancestral stock. Of late years,
however, the latter species has grown greatly in favour for the
distinction, partly on account of the unquestioned adoption and
reiterated publication on the part of recent writers of a statement
made many years ago, but none the less erroneous, that the
Domestic Cat has a longer and more tapering tail than the Wild
Cat of Europe*, and partly on account of Nehring’s assertion 7 that
the Domestic and Egyptian Cats resemble each other, and differ
* Macgillivray, in his excellent account of the British Wild Cat (‘British
Quadrupeds,’ Jardine’s Naturalist’s Library, pp. 188-195, 1854), long ago pointed
out that the tail of the Domestic Cat is thinner and more tapering than that of the
Wild Cat, because its fur is much shorter. Dr. Hamilton also exposes the fallacy of
the above-stated belief (‘ Wild Cat of Europe,’ pp. 41-42, 1896). One modern writer
speaks of the tail of the Wild Cat as “clubbed.” I have never seen a specimen with
a tail to which this epithet meaning broader at the tip than at the base could be
applied.
+ SB. Ges. nat. Freunde Berlin, 1887, pp. 26-27, & ‘ Humboldt,’ 1888, pp. 189-141.
Nehring (Zeits. fiir Ethnologie, xxi. pp. 558-9, 1889) suggests that Domestic Cats
are descended from two main stocks—one from S.E. Asia, the other from N.E.
Africa. From the former arose the ‘Chinese? Domestic Cat; from the latter the
Egyptian House-Cat. The Egyptian House-Cat was the forerunner of the European
Domestic Cat, with an infusion, especially in Germany, of the European Wild Cat.
But whether the stock of the Asiatic House-Cat was F’. inconspicua Gray, or
F. manul, or some other smaller Asiatic species, the author leaves undecided. It does
not appear that Nehring realised the existence of the two types of European Domestic
Cats I have described. Also he can scarcely have been acquainted with the
peculiarities of F'. manul or with the nature of the affinity between F. incon-
spicua and F. ucreata.
156 MR. R. I. POCOCK ON [Feb. 19,
from the Wild Cat in having the back of the hind Jeg from the hock
to the pad blackish. This statement is negatived by three facts :—
the area in question is not blackish in all Domestic Cats; nor in
all examples of the Egyptian Cat; nor does it lack the dark tint in
any specimens of the Kuropean Wild Cat that I have examined.
The prevalent idea on this subject has been expressed as follows :—
“The black sole of the foot suggests that the Caffre Cat is the
chief stock from which the Domestic Cats of Kurope have been
derived”; and “that the Kuropean Wild Cat was not the direct
descendant |? lapsus for ancestor] of the domesticated breeds of
the western part of the continent is rendered pretty evident by its
short and clubbed tail, to say nothing of the absence of dark soles
to the hind feet” *. And again:—‘It has been maintained
by many naturalists that the European Domestic Cat is chiefly
derived from the north-eastern race of this species [/’. ocreata]
found in Egypt; at least, the domestic form is certainly not
derived from the European Wild Cat” +. The foundation for this
positive opinion entertained by Mr. Sclater may be found, I
think, in his belief (Cat. Mamm. Indian Mus. ii. p. 934, 1891),
taken from Blasius, that in the skulls of the Domestic Cat the
frontal and squamosal bones are separated from one another by
the parietals and alisphenoids, and the nasals are not produced
posteriorly beyond the frontal processes of the maxille, the con-
verse being the case in the Wild Cat. I have before me the |
skulls of four ‘ London” Cats. In one of them the nasals project
well beyond the maxille, in two as far back as the maxille, and in
the fourth not so far. Again, the distance between the frontal and
the squamosal is in one skull as much as 3 mm., in two others it is
only about 1 mm. or less, while in the last the squamosal and
frontal touch on one side but not quite on the other. Nevertheless,
it is perfectly true that, as a very general rule, the squamosal
and frontal are separated by the junction of the parietal and ali-
sphenoid in Domestic Cats as they also arein J’. ocreata. But
this is also the case in both the skulls of the Scotch Wild Cat that
I possess=. Im one of them the parieto-alisphenoid bridge
measures 4 mm., in the other 2mm. The bridge is also present
in one of two skulls of this species in the Museum of the College
of Surgeons, whereas in the other the frontal and squamosal have
a long sutural union.
Thus, as Dr. Hamilton has already shown, the claims of both
Blasius and Nehring regarding the differences above alluded to
between the Wild and Domestic Cats of Europe will not stand
the test of the examination of skulls and skins.
It appears to me that much barren discussion on this subject
would have been saved by the realisation of the closeness of the
affinity between the Egyptian and the European Wild Cats
* “Cats &.’ in Allen’s Nat. Library, pp. 156 & 157 (tee), by R. Lydekker.
+ ‘Mammals of S. Africa,’ i. pp. 43-44 (1900), by W. L. Sclater.
{~ There is no reason to doubt that both these skulls belonged to pure-bred Wil
Cats; this I can vouch for, since I have the skins.
1907. | ENGLISH DOMESTIC CATS. 157
(Ff. ocreata and F. sylvestris). The type of pattern they present
is not found in any other species of Felis, the nearest approach
thereto being that of the Tiger (/. tigris) and in a remoter degree
of Pallas’s Cat (7. manwl). The similarity in pattern between the
two, coupled with their geographical distribution, almost induces
the adoption of the view that they are but northern and southern
forms of the same species. There is as yet, however, no positive
evidence that they actually intergrade to the extent of justifying
the conclusion that their distinctive features are merely of sub-
specific value.
The characters they have in common are as follows :—
Close resemblance in the shape, size, and structure of the skull
and teeth *.
General similarity in size and shape of the body, of the paws
and ears,and in the length of the tail, though this organ is
apparently a little longer at least in North-African examples of
F. ocreata than in F. sylvestris.
General similarity in pattern: in both species the sides of the
body are typically marked with wavy vertical dark stripes ex-
tending from the spine to the belly, and better defined, as a rule,
on the thoracic than on the abdominal portion ; the upper parts
of both fore and hind limbs are transversely banded with broad
bars, almost always darker than the transverse stripes on the
body ; the fore leg is black below from the toes to the wrist and
behind at the elbow, the two internal brachial stripes, normal in
felines, being well developed ; the hind leg is normally blackish
below from the toes up to the hock. The distal portion of the
tail has a black tip and about three well-defined black stripes pre-
ceding it, the proximal portion of the tail being much less distinctly
barred and marked at best with an ill-defined median dorsal
stripe. The throat is unstriped or only indistinctly striped, and
usually exhibits a white spot ; the thoracic and anterior abdominal
areas of the ventral surface are spotted ; the posterior abdominal
and inguinal areas are unspotted and tinged with yellowish buff,
especially on the inner side of the thighs. Identity prevails even
in the colour of the individual hairs, which are slate-grey at the
base, then buff or cinnamon, then black, the distal portion being
yellowish or greyish white with a black tip.
The principal differences between them are as follows :—
in F. sylvestris the four paired stripes on the head and neck are
well defined; on the occipital region they diverge from the middle
line and run backwards almost to the shoulder as four wavy widely
* Amongst the skulls in my own collection and in that of the British Museum I
have so far failed to find any constant characters for distinguishing the two species.
By the sum of a number of small features in the teeth and bones, the skull of
sylvestris can generally be recognised from that of ocreata. But up to the present
every character which I thought might prove to be distinctive of sylvestris has
broken down under the examination of a long series of skulls of ocreata, a species
‘which, in its broad sense, is extremely variable in its cranial osteology. So far as the
skull is concerned, the differences between the two species must be regarded as of
“ subspecific ” value.
158 MR. R. I. POCOCK ON [ Feb. 19,
separated stripes, the admedian pair being, as a rule at all events,
better emphasised than the laterals, which lie quite at the sides of
the upper surface of the neck. In F. ocreata the head- and neck-
stripes are usually badly defined ; when present on the neck they
are narrow and lie close together.
In F. sylvestris there is generally a very distinct black wavy
median spinal stripe, usually extending from behind the shoulders
to the root of the tail. In /. ocreata the entire spinal area is
markedly darker than the sides of the body, sometimes showing
traces of three narrow stripes; but the median is never so strong
as in J. sylvestris. In the latter, however, a pair of narrow
interrupted latero-dorsal stripes is sometimes traceable.
These are the most obvious distinctions. Others are less
constant. For instance, in /. sylvestris the ears are, generally
speaking, of the same colour as the head, though not infrequently
they are washed with yellow either all over or only towards the
tip. In /. ocreata they are almost always yellower or redder,
generally very decidedly so, especially in African specimens ; but,
on the other hand, in an example of /. ocreata sarda in the British
Museum the yellow on the ear is no more conspicuous than it 1s
in some examples of /”. sylvestris.
In F. sylvestris the coat is ionger and thicker than in F. ocreata.
This imparts to the former Cat a heavier, more robust, and
shorter-legged appearance, and especially suggests that the tail
is blanter at the apex *. The value of this difference is discounted
by the fact that /. sylvestris is a more northern type than
F. ocreatat, and that the length and density of the fur varies a
good deal in the latter, which appears to be a species endowed
with great capacity for environmental adaptation both as regards
coat and colour.
Yet, in spite of the obvious resemblances above mentioned, the
assumption of the total diversity of the two forms seems to have
been pretty general, if we may judge by the absence of all com-
arison between them in monographs of the Felide.
Now the characteristics which the Egyptian and European Wild
Cats have in common are all possessed by the Domestic Cats of
the “striped” type; and they are not found in any other species
of Felis known to me. Hence there is no difficulty in the way of
believing that our “striped” Cat is the direct and but little modified
descendant of either /’. sylvestris or Ff’. ocreata, or probably of both
combined. /. sylvestris inhabits Spain, Italy, Greece, and Asia
Minor; and /. ocreata Egypt, Tunisia, Algeria, and Sardinia.
Thus one species or the other is found in the countries bordering
the Mediterranean basin, where the civilisations of EKurope had
their origin. If Egyptian Cats were taken to Greece, Italy, or
* Quite similar differences in the thickness of the tail may be seen in Siberian
and Indian Tigers.
+ A very good description and an excellent figure of this species may be found
under the name Felis lybica in Anderson’s and de Winton’s ‘ Mammals of Egypt,’
pp. 171-176, pl. xxiv. (1902).
1907. | ENGLISH DOMESTIC CATS. 159
Spain, it is highly probable that they interbred with the native
Wild Cat, especially at a time when the latter was far more
abundant than it is now and when precautions to prevent the
tame animals from straying are not likely to have been rigidly
observed. But even if such crossing took place, I do not believe
its effects could be traced in the progeny with any certainty, on
account of the resemblances between the parent forms.
It may however, in my opinion, be assumed that the differences
our English striped Domestic Cats exhibit from /’. sylvestris on
the one hand and from /’. ocreata on the other, coupled with an
unmistakable likeness to both, are attributable to that cause. To
assume that the striped pattern of this Cat is due to interbreeding
between domestic “ Tabbies” or Blotched Cats and specimens of
F, sylvestris, needlessly complicates a quite simple question.
It is difficult to decide which of the two species our Striped
Cats most resemble. Typical short-haired individuals recall
fF. ocreata in the length of the fur on the body and tail; whereas
in the colour of the ears greater similarity is presented to
F. sylvestris than at all events to African examples of /’, ocreata.
In the proximity of the stripes on the neck, resemblance is evinced
to F. ocreata ; but their distinctness recalls, though in a lesser
degree, those of /. sylvestris. In the distinctness of the spinal
stripe, the domestic form les nearly midway between the two;
but the transverse stripes on the body and tail are as a rule more
sharply defined than in typical members of either of the wild
species: but their want of definition in some long-haired specimens
is quite paralleled by that of /. sylvestris, and suggests that their
indistinctness in the latter is attributable to length of fur.
In the length of the tail the Striped Cat seems to be nearer
F, sylvestris. There is no evidence that this organ is shorter in
the domestic form than in the wild species just mentioned ;
whereas in North-African examples of /. ocreata the tail is
longer than in either of those types*.
In connection with the likeness this Cat presents both to Felis
ocreata and F’, sylvestris, it is apposite to note that out of four
authors who have described specimens under the belief that they
represented wild species or races, two compared them to F’. sylvestris
and two to #. ocreata. Blyth tT, speaking of an example from the
Punjab Salt Range, says “itis of the streaked or spotted type,
the colouring and markings of which are not much unlike those
of the European Wild Cat (/. sylvestris)” ; and Mivart {, in his
description of /. torquata, remarked, “this Cat has much
resemblance to the European Wild Cat.” Blanford, on the
contrary, says “the characters of the upper premolars distinguish
F. torquata from the allied /. caffra (or caligata) | =ocreata]” § ;
* Anderson and de Winton, ‘Mammals of Egypt,’ P- 172.
+ J. A. S. Beng. xxv. p. 442 (1856). t ‘The Cat,’ p. 420 (1881).
§ ‘Mammals of British India,’ p. 86 (1888). The character Blanford refers to is the
nearness of the first maxillary premolar to the second—a very variable feature in the
skulls of Domestic Cats and also, though in a lesser degree, in those of F’. ocreata.
160 MR. R. I. POCOCK ON [ Feb. 19,
and Miss Bate described a specimen from Crete as the type of a
distinct subspecies of /. ocreata *.
The origin of F’. catws appears to be at present quite unknown.
Tt may be held :—
a. That it arose per saltwm as a sport from the other Domestic
Cat (torquata), and that the pattern persisted im virtue of
its own inherent dominance without the aid of Man, or in
virtue of the guiding factor of selective breeding. In
opposition to this must be urged the complete absence
of evidence that species of elis are ever dimorphic in
pattern, and the ascertained fact that they breed true to
the specific or subspecific type.
b. That it arose from torqguata by the slow and gradual process
of preserving and breeding from fancied varieties. But in
answer to this it may be pointed out that there is no reason
to think that selective breeding of Cats was ever seriously
practised until the latter portion of the nineteenth century.
Moreover, if the catus type arose by that process, inter-
mediates between it and torguata would probably be seen
everywhere.
c. That it resulted from the interbreeding of /’. ocreata and
F. sylvestris. When two distinct species cross, the hybrid
sometimes reverts in some respects to the characters of a
common ancestor of both. There is no reason, however,
for thinking that the pattern of catus was the pattern of
the ancestor of sylvestris and ocreata; and it seems to be
in the highest degree improbable that the progeny of two
closely allied and similarly striped species like sylvestris
and ocreata should be marked in a totally different manner
from its parents. There is, moreover, good reason for
thinking that the torqguata-breed was the resulting hybrid
of that cross.
d. That it resulted from interbreeding between sylvestris or
ocreata or torquata and some exotic species introduced into
Europe. ‘There is, however, no reason to believe that either
tamed or wild representatives of any exotic species other
than ocreata were so introduced, apart from menagerie- kept
animals.
e. That it is the direct descendant of some existing exotic species.
It is quite evident, however, that it is not the direct un-
modified descendant of any known species of Felis, since its
pattern is unique in just the same way and to the same
extent that the pattern of the Tiger is unique in the genus.
f. That it is the survivor of some extinct, probably Pleistocene
Cat of Western Europe. By the method of exhaustion of
other possibilities, one falls back upon this supposition,
which at least has this in its favour, that no very obvious
or cogent reason can be advanced against it.
* P. ZS. 1906, p. 317.
1907.] ENGLISH DOMESTIC CATS. 161
4. On Manz, Persian, Siamese, Indian, and Abyssinian Breeds.
As already explained, ‘‘ Manx” Cats may be either of the catus
or torquata type. Apart from the abbreviated tail, which has
been discussed and dismissed as of no systematic significance from
the zoological standpoint, Manx Cats differ or are alleged to differ
from ordinary short-haired Cats in standing relatively higher at
the hind-quarters. I have not seen any measurements sub-
stantiating this claim ; and it is dificult to decide to what extent
the greater apparent posterior stature is an optical illusion caused
by the absence of the tail. It is quite possible, however, that
suppression of the tail is correlated with greater height at the
sacrum, and that, to put it crudely, the caudal material is dis-
tributed to or absorbed by the hind-quarters. Some Lynxes
certainly seem to stand higher on the hind legs than the majority
of species of Felis. Be thisasit may, the circumstance that height
at the posterior region is considered by fanciers ‘“‘ a point” in the
Manx breed throws the fact—if fact it be—under suspicion of
having been fostered by selective breeding and of being therefore
unworthy of consideration from the phylogenetic and systematic
point of view, zoologically speaking. In other words, as little
importance should be attached to the character as to the absence
or abbreviation of the tail. .
With regard to so-called ‘“‘ Persian ” or ‘* Angora” Cats*, there
seems to me to be no reason to suppose that any other species is
involved in their ancestry than in the ancestry of the short-
haired breeds most common in Europe. Both the “blotched”
and the ‘striped ” styles of pattern occur; and no other type of
pattern is known amongst them, so far as I am aware. The
skull, moreover, is not distinguishable from that of short-haired
Domestic Cats. The small systematic value that should be
attached to the long coat has been already insisted upon. Nothing
seems more likely than that tame Cats were imported from
Egypt into Persia; the European Wild Cat (7. sylvestris) occurs _
both in Asia Minor and Persia, and F. ocreata has been recorded
from Syria and Arabia. Hence it seems needless to look beyond
the two species just mentioned for the origin of “striped ”
** Persian” Cats, even on the supposition that they came ori-
ginally from Persia. That the ‘‘ blotched ” Persians had the same
origin, whatever that may have been, as the“ blotched” short-haired
Cats is in the highest degree probable. For myself, I think
it quite needless to consider that so trivial a character as long
hair is to be traced in all cases to Cats imported into Western
Europe from Asia Minor or Persia. The suggestion—first made,
I believe, by Pallas, but repeated even in modern literature on
the subject—that the Central Asiatic species, Pallas’s Cat (Felis
manul), contributed to the “ Persian” breed, has nothing to be said
in its favour. Felis manul is quite unlike all domestic breeds.
* Desmarest (Nouv. Dict. vi. p. 122, 1816) gives Anatolia as the original country
of this breed.
Proc. Zoot. Soc.—1907, No. XI. 1]
162 MR. R. I. POCOCK ON [ Feb. 19,
The ears are small and set very low on the sides of the head,
leaving a great width of forehead between them; the facial and
body markings are different; and, lastly, the pupil of the eye
contracts to a circular disk *.
There is, however, one species of Felis which must not be
altogether forgotten in considering the possible origin of Persian
Cats. This is the Bokhara Steppe Cat (/elis caudata), described
by J. E. Gray (P. Z.8. 1874, pp. 31-32, pls. vi. & vii.). The type
is in the British Museum. ‘The figures of the entire animal and
of the skull show that this Cat is closely related to Felis sylvestris
and to /. ocreata, and not to /. chaus, with which Dr. Gray com-
pared it. It has a thick coat and bushy tail, like 2. sylvestris ;
but there are no distinct spinal stripe nor definite stripes on the
head and neck. In length the tail resembles that of /. ocreata ;
and likeness to this species is further shown by the indistinctness
of the head, neck, and dorsal stripes. From both the other species
it differs in being covered with small spots; but these spots, at
least on the anterior part of the body, show, I think, signs of the
coalescence into transverse rows which is realised in /’. sylvestris
and /. ocreata. Gray describes the colour as yellowish. The
skin, which is probably faded, might be more aptly described as
‘““ounce”-grey. Whether or not this species has any connection
with the “spotted” Cats of the Punjab Salt Range, mentioned
below (p. 164), I am unable to say.
It is worth putting on record the fact that in two out of the
three skulls of “ Persian Cats” I possess the jaws are slightly
“underhung,” that is to say, the mandible protrudes a little in
front of the premaxillz. I have not noticed this peculiarity im the
skulls of any other Cats, either wild or domestic. It may be purely
accidental, or it may indicate that the taste of fanciers in Cats is
running along the same lines as those of breeders of Domestic Dogs.
In this case we may in the future have a race of snub-nosed Cats
departing in facial elegance from Nature’s type of Felis in the
same manner that Pugs and Bulldogs depart from Nature’s type
of Canis.
The origin of the Siamese breed has been a much-discussed
puzzle. The peculiar coloration fT must be set on one side as value-
less towards affording a clue. The cats are obviously albescent, as
is attested by the hue of the hair and frequently by the blueness
of the eyet. But the albescence, complete in the newly-born
kitten, is mitigated as age advances by a melanistic taint which
evinces itself in two ways, both of unusual occurrence in Nature.
In the first place, the black pigment is distributed symmetrically,
and primarily affects the face and ears, the tail, and the legs.
* The illustration of this species in Elliot’s ‘ Monograph of the Felid,’ pl. x., was
taken from a stuffed specimen, and does not correctly depict the peculiarities of the
head and eyes. Elliot, however, very rightly insists that this Cat is unlike all other
known species.
+ Mr. E. G. B. Meade Waldo has reminded me that the so-called Himalayan breed
of domestic rabbits is almost identically coloured.
+ R. I. Pocock, Ann. Mag. Nat. Hist. (7) xix. p. 194, 1907.
ISO] ENGLISH DOMESTIC CATS. 163
In the second place, it increases in quantity as age advances, and
gradually encroaches upon the whiter areas, converting the hair
of the body from whitish or pale fawn into pale or dark brown.
Sometimes there are indistinct traces of spots on the hind-quarters
and legs.
Judging from their size, form, and general aspect, I should say
these Cats are nothing but a domestic variety of /. ocreata. The
only skull of this breed that I possess * is that of a female with its
contours and ridges indicating considerable muscular development.
It is a short, broad skull, with expanded zygomata, and beyond all
possibility of doubt falls into the group typified by /. ocreata and
F’. sylvestris. In profile it is noticeable that the highest point of
the cranium lies well behind the fronto-parietal suture, and that
the area in front of that point, almost as far as the nasals, is nearly
flat and slightly inclined. This is not a common feature in the
skulls of Domestic Cats; but it is almost exactly paralleled in the
skull of an old male specimen of /. screata from Suakin, and does
not occur in the skull of a Siamese Cat in the British Museum,
in which the highest point of the skull is on the frontal bones,
which are evidently swolien just behind the postorbital processes.
Another peculiarity in my specimen is the division of the infra-
orbital foramen by a bridge of bone. I find this feature repeated,
however, in the skull of a London Cat not referable to the Siamese
breed ; and it does not occur in the Siamese skull in the British
Museum.
Nevertheless, these two Siamese skulls agree in the possession
of two small characters, one of which I can only match in one of
the English Cats’ skulls I possess, while the other cannot be quite
matched in any of them. The first character is the height of the
interparietal crest, which is better developed than is usually the
case in English Domestic Cats, though it is equalled in the skull
of one London Cat that I have seen, and of one feral example of
the torquata-type of F. ocreata, from Celebes, in the British
Museum. The second character is the greater narrowness of the
posterior portion of the nasal bones and the more marked abrupt-
ness of the constriction between this portion and the expanded
anterior portion. But since the two Siamese skulls differ in
degree with respect to this character, and since the nasal bones
vary greatly in length and width in English Domestic Cats, no great
importance can, I think, be attached to the feature in question.
Moreover, as already stated (p. 152), the Celebes example of
torquata in the British Museum also has the nasals compressed.
Since, therefore, the colour of the Siamese Cat affords no
evidence of its descent, and the skull is decidedly of the ocreata+
sylvestris type, there seems to me to be no reason to look beyond
those species or indeed beyond ocreata for its origin. Those who
claim that it has had an origin independent of our own Domestic
* Tam indebted to Mr. F. W. Cousens, F.Z.S., for the skin and skull of one of
the two specimens of this breed I have been able to examine. Mr. Cousens also
kindly gave me the skins and skulls of two Persian Cats.
Iie
164 MR. R. I. POCOCK ON [ Feb. 19,
Cats from some Siamese or Oriental species, must be challenged
to produce that species before the question can be profitably
discussed. There seems to be no reason whatever for entertaining
Trouessart’s suggestion* that the rare Bornean Felis badia was
its agriotype. I may add that all the small species of Felis in-
habiting Siam, including even /. chaus, which is doubtfully
indigenous, differ in the structure of the skull and teeth from
the Siamese Domestic Cat 7.
According to Blyth (Journ. As. Soc. Bengal, xxv. pp. 442-445,
1856) two types of Domestic Cat were prevalent in India in his
time. ‘One is the streaked or spotted type, the colouring and
markings of which are not much unlike those of the Huropean
Wild Cat (Ff. sylvestris), only more distinct, and the transverse
streaks are more broken into spots, especially towards the hinder
part of the body ; the fur, however, is short, and the tail £ slender
and of uniform apparent thickness to the end, showing a series of
rings and a black tip; ears slightly rufescent externally but
infuscated, but passing to black at tip, where there is a distinct
small pencil-tuft of black hairs. Paws deep sooty black under-
neath.” Two examples of this Cat were shot wild in the Punjab
Salt Range; another seen at Allahabad in a state of domestica-
tion exactly resembled them ; but the tame Cats of Calcutta were
usually greyer, with smaller and more numerous spots.
The other type, says Blyth, resembles the Jungle-Cat (/. chaus)
in colour, the body being uniformly “ cat-grey,” more or less rusty
or fulvescent, without a trace of spots or stripes, but the stripes
on the tail and limbs are more distinct; there are also confused
stripes on the forehead and two on the cheeks and a dark band
across the chest; the lower parts are whitish or tinged with
tawny, and spotted; the ears are dull rufous behind with a
slight blackish tip and no pencil of hairs; the paws more or less
sooty beneath. In its proportions, however, this Cat is quite
different from /’. chaus, the limbs and ears being much shorter,
and the tail much longer and tapering. Blyth adds that Domestic
Cats of this type abounded in Bengal, if not all over India, but
were quite unknown in Europe; whereas, on the other hand,
the English “tabby” was quite unknown in India.
Mr. W. L. Sclater also discussed these two breeds, and accredited
Blyth with the belief that the self-coloured ‘“ chaus”-like type was
derived from interbreeding between the ‘‘ Domestic” Cat and
F. chaus (Cat. Mamm. Ind. Mus. 11. p. 233, 1891). He also sug-
gested that the form named /. torquata by F. Cuvier was based
upon a feral example of the spotted or streaked breed.
So far as I can judge from what the two authors quoted say
about these breeds, there is nothing to distinguish them from
* Cat. Mamm. 1904, p. 273.
+ For list of the species, see S. S. Flower, P. Z.S. 1900, pp. 322-327.
~ In this particular the tail must resemble and not differ from that typical of
F. sylvestris. Surely Blyth intended to say “tapering” towards the end, otherwise
the remark is pointless.
1907. | ENGLISH DOMESTIC CATS, 165
F’. ocreata, which may be either self-coloured like the ‘ chaus”-
like Cat or spotted and streaked like the spotted Indian Cat. The
facts adduced do not appear to me to supply any evidence that such
indigenous Indian Cats as F’. rubiginosa, PF. chaus, or F’. benga-
lensis have contributed tothe strains; and I strongly suspect that
they have been derived from /, ocreata either by the importation
of tamed specimens or by the reclaiming from the wild state of
examples of this species which may have inhabited India in com-
paratively recent times.
In the British Museum there are a few skins of Indian
Domestic or semi-domestic Cats belonging to the spotted or
striped and to the self-coloured types mentioned by Blyth. An
examination of them fully confirms the opinion I had formed
from reading Blyth’s remarks. Those belonging to the striped or
spotted type are, as Mr. Sclater suggested, the same as the form
described by Cuvier as torguata. ‘Those of the self-coloured type
do not differ from them more than some of the self-coloured Cats
differ from the striped Cats to be seen in the London streets.
Some of them are more rufescent than others; but I cannot find
in them a particle of evidence of partial descent from 7. chaus
or from any other indigenous Indian species.
Not uncommonly Indian Domestic Cats differ from typical
English examples of torquata in having the bands on the tail
narrower and the stripes, especially on the forehead and cheeks,
more rufous. The narrowness of the caudal stripes arises from
the splitting of the normal stripes; but this character as well as
the rufescence may be seen in African examples of /. ocreata.
There is no reason, therefore, to doubt that the Indian Domestic
Cats are descended from that species. It must, of course, be con-
ceded that they may interbreed with indigenous Indian species,
and especially with the so-called Desert-Cat (2. ornata), which,
from the structure of the skull and teeth, as well as from other
characters, must be regarded, I think, as the Indian representa-
tive, as /”. caudata is the Bokhara representative, of the group
typified by /. ocreata in Africa, and by F’. sylvestris in Kurope.
Cats of the so-called “ Abyssinian” breed may be descended, for
anything I know to the contrary, from specimens of 7. ocreata
directly exported from Abyssinia. They are certainly not unlike
some self-coloured examples of that species. On the other hand,
it would I imagine be difficult to separate them from fulvescent
“ Ticked” Cats, which appear to me to be nothing but examples
of the torquata-type in which the pattern is broken up and
evanescent (see p. 152).
On alleged Cases of Interbreeding between Domestic Cats and
various wild Species of the Genus Felis.
It has been stated over and over again that Domestic Cats
interbreed freely with the native Cats of various species inhabiting
the countries to which they have been transported. One cannot
166 MR. R. I. POCOCK ON [ Feb. 19,
help wishing there was more positive evidence of the fact. It
may be so; but the statements of authors on this subject cannot
be accepted as of any great value unless there is collateral evidence
of their intimate acquaintance with the wild species in question
and with the range of variation in colour, pattern, and structure
of the domestic breeds. Many alleged cases of interbreeding may
be found quoted in Darwin’s ‘ Variation of Animals and Plants
under Domestication.’ It is very likely that some Cats seen
by Jardine in the north of Scotland were rightly regarded as
hybrids between domestic animals and J. sylvestris. And
St. Hilaire’s statement that the Domestic Cats of Algiers cross
with the native Cat (/. lybica = ocreata), and Layard’s to the
effect that similar crosses occur with /. ocreata caffra in
S. Africa, may also be true. But in the last two cases it may be
doubted if evidence of the crossing would be shown by the progeny ;
and as regards Jardine’s alleged hybrids, long-coated specimens of
the “striped” Domestic Cat might easily be mistaken for half-
bred Wild Cats. A striped Cat known to be descended from a
feral specimen was recently caught in the New Forest, where
Felis sylvestris has not occurred for at least a century, and sent to
the British Museum by Mr. P. H. Barker as a hybrid. On the
strength of its long coat, it must be regarded as a ‘“ half Persian.”
Had it come from parts of the north of Scotland, where /. sylvestris
still lingers, the entire purity of its descent might have remained
for ever in doubt. As regards the claim that Domestic Cats in
India interbreed with the Jungle-Cat (/’. chaus), I am unable to
find evidence that satisfies me of the certainty of this occurrence
(cf. supra, p. 165). In one case the claim is actually based, in
part at all events, upon the occurrence of the internal brachial
stripe on the fore leg, a feature regarded by the observer as
characteristic of F’. chaws, whereas it is the most persistent of all
the stripes in the genus els.
No one would be so rash as to atlirm that interbreeding does
not occur between Domestic Cats and even widely different wild
species. But unless the skins and skulls of alleged hybrids are
forthcoming for examination, there is no basis for the discussion
of the question. Vague suppositions of observers cannot be
regarded as evidence of the fact; and I am of opinion that the
prevalent beliefs on the subject are to be assigned in a great
measure to the observation of semi-wild specimens of the common
but little known striped Domestic Cat, which may be either red
or grey in colour, and either striped or spotted in pattern.
6. Summary.
The substance of the foregoing remarks may be epitomised as
follows :—
1. The characters used by breeders and fanciers as a basis for
their so-called breeds of English Domestic Cats have no
scientific value, in the sense of affording a clue to affinity
and descent.
1907. | ENGLISH DOMESTIC CATS. 167
i)
. The pattern—or, in other words, the arrangement of the
stripes—shows that English Domestic Cats are referable
to two distinct types, whether they belong to the ‘‘ Manx,”
‘“« Persian,” or ‘ Short-haired ” breeds.
3. These two types of pattern are different in kind and do not
intergrade. They are so distinct from each other that no
one would hesitate to regard them as characterising two
well-marked species if the animals presenting them existed
in a wild as opposed to a domesticated state.
4, In one type of pattern the stripes take the form of narrow
transverse or vertical bands which sometimes break up
into spots. To feral or domesticated examples of this Cat
have been given many names, of which torquata is the
best known and angorensis or striata possibly the oldest.
5. This Cat (torquata) was apparently domesticated in Europe
at least as early as the 16th century. There seems to be
no reason therefore for regarding it as of Indian origin.
6. It closely resembles in pattern two existing species, namely,
the so-called Egyptian Cat (/. ocreata) and the European
Wild Cat (/. sylvestris), both of which occur at the
present day in the Mediterranean Region, and are very
nearly related to each other. There is no difficulty in
the way of believing that they are the ancestral forms or
‘“‘aoriotypes” of this domesticated race (torquata).
7. In the other type of pattern the stripes take the form of
broad longitudinal or obliquely longitudinal bands forming
a ring-like or spiral arrangement on the sides of the abdo-
men. To domesticated examples of this Cat, Linneus gave
the name catus, which cannot be applied to any other form
of the genus Felis. Domestica is its best-known synonym.
8. This Cat (catus) is certainly known to have been domesti-
cated in Europe in the middle of the 18th century. It
was not, however, apparently known in India in the
middle of the 19th century. Probably, therefore, it is of
European descent.
9. Its origin is unknown. Of the several hypotheses that may
be held on this subject perhaps the following two are the
most to be commended :—that it arose as a sudden varia-
tion or sport from the forquata-breed, in which case
European Domestic Cats are dimorphic in pattern; that 16
is the direct descendant of some extinct Pleistocene Cat,
in which case there are two distinct species of Domestic
Cat in Europe.
EXPLANATION OF THE PLATES.
Puate VIII.
Blotched Tabbies, Felis catus.
The four skins photographed were selected out of a large number to show the
principal variations to which the pattern in this Cat is liable. All came from the
Cats’ Home in Camden Town, London.
168 DR. C. G. SELIGMANN ON DEATHS [Feb. 19,
Prats IX.
. Striped Tabby, Felis torquata.
Fig. 1. Example of the Felis sylvestris-type to be compared with fig. 1, Plate X.
2. Like the last, but the pattern more spotted.
3. A partially albino specimen with the spots larger and more widely spaced
than in No. 2.
4. Specimen of the so-called “Ticked ” breed, with the spots disintegrated and
generally distributed over the body.
(From the Cats’ Home, Camden Town.)
to}
Puate X.
Agriotypes of the Striped Tabby, 7’. torquata.
Figs. 1 & 2. European Wild Cats, F. sylvestris, from Ross-shire in Scotland.
Figs. 3, 4, 5. Young examples of the South African race of the African Wild Cat,
Felis ocreata caffra. The pattern is usually more distinct in the young
than in the adults of this species. The exact lscality of these three
specimens is unknown. They were shipped from Cape Colony.
(The camera has emphasised the pattern of the skins depicted on this Plate.)
Report on Deaths occurring in the Society’s Menagerie
during 1906. By C. G. Seniemany, M.D., F.Z.8.,
Pathologist to the Society.
[Received February 19, 1907. |
(Text-figure 61.)
In the annexed table will be found the causes of death so far as
they could be discovered of 355 mammals and 283 birds which
died in the Society’s Gardens, and which were submitted to post-
mortem examination during the year 1906. In these mammals
and birds no cause of death is stated to have been found in 36
mammals and only one bird, but 1t must be noted that the number
of deaths put down to trauma and exhaustion is much larger than
in 1905. This is due, in my opinion, to the important part
played by depressed vitality, and in some cases darkness and
cold, in bringing about death. The method of classification is the
same as that adopted last year*, but two new headings occur, viz.
Diseases of the Ductless Glands and Deaths due to Old Age.
Possibly two of the deaths among birds attributed to “ trauma
and exhaustion” really took place owing to the effect of parasites.
The following remarks refer to conditions of special pathological
interest occurring in the animals in the Gardens.
Tuberculosis. ae is yet too early to attempt any full appreciation
of the effect of thoroughly disinfecting, scraping, and repainting
the monkey-house, undertaken early in the year, when the heating
arrangements also were altered, but the diminution of deaths from
tuberculosis—34 in 1906 against 53 in 1905—is decidedly
encouraging. If the figures be expressed as percentages, 21°6 of
the total deaths in monkeys were from tuberculosis in 1906 against
* “Note on Deaths occurring in the Society’s Gardens during 1905,” P. Z.S. 1906
p. 234 et seq.
169
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170 DR. C. G. SELIGMANN ON DEATHS [ Feb. 19,
35°8 in 1905. Assuming that there were about 100 monkeys in
the house on January the Ist, 1906, between 16 and 17 per cent.
of the inmates of the house have died of tuberculosis during the
past year, since the actual number of monkeys received was 105.
My thanks are due to Mr. Pocock for ascertaming the number
of admissions during the year, and I am further indebted to him
for the estimate of 100 as the number of monkeys in the house
in January 1906.
Much attention has been paid to tuberculosis in birds, and
certain interesting conclusions concerning avian tuberculosis can
be drawn from a study of the material available. It must in the
first place be noted that during the past two years about 30 per
cent. of all deaths in birds have been due to tuberculosis; and the
more the matter is Investigated the more obvious it becomes that
a very large percentage of the cases of tuberculosis occurring in
birds in the Zoological Gardens are the result of infection by the
intestine, which can hardly be due to any other cause than the
swallowing of particles of contaminated soil while food is being
picked up. By far the greater proportion of birds dying of
tuberculosis in the Gardens present typical lesions in their spleen
and liver, which can only be explained on the hypothesis of an
ingestion tuberculosis: sometimes the intestine is affected, but
more often this is not the case, and typical tubercular ulceration
of the gut in birds dying in the Gardens is rare. But although
ulceration of the gut does not frequently take place, enlarged
tuberculous glands at the root of the mesentery are by no means
uncommon; a condition akin to tabes mesenterica of human
pathology being set up without obvious damage to the mucous
membrane of the gut, but with the addition of lesions in the
spleen or liver or both. Even where the intestine is affected,
ulceration is rare or slight; comparatively large submucosal
nodules being formed over which the villi of the mucous mem-
brane often seem enlarged, so that a curious condition suggestive
of multiple closely-set warts is found to occupy the inner surface of
the bowel. Sometimes, as in a Vulturine Guinea-fowl (Acryllium
vulturinum), the whole of the large gut may be thickly studded
with these warty growths, which on section are found to
contain dense masses of acid-fast bacilli. In other cases the
whole of the small gut is similarly affected, as in a Burmese Slaty-
headed Parakeet (Palecornis sp.). In the latter case it was inter-
esting to note that there were no lesions in any other abdominal
organ or in the thorax, while in the case of the Guinea-fowl there
was early tuberculosis of the lung, liver, and spleen.
Pneumonia and Broncho-pneumonia._—No marked improvement
in these diseases has followed the cleaning of the monkey-house.
Mycosis.— Reference was made last year to the occurrence of a
disease, mycosis, which was due to the invasion of the tissues by
a mould, Aspergillus fumigatus. This disease is by far commoner
in water-fowl than in other birds, and when attacking these its
characteristic lesions are usually widely distributed throughout
the body-cavity ; but a case occurred in an African Buzzard (Buteo
1907. ] IN THE SOCIETY'S MENAGERIE. el
desertorum) in which the disease, obviously not very recent, affected
only the air-sac and the lung of the right side, there having been
no extension of the parasite by continuity or by infection of the
blood or lymph stream.
Cardiac Failure —It has been noted in the case of Ostriches,
Rheas, and Cassowaries, as well as some of the larger Storks kept
Text-fig. 61.
Aorta of Tiger, showing several aneurysms.
in the Gardens, that their hearts after death are usually extremely
flabby, while the subpericardial fat may be replaced by a loose-
172 DR. C. G. SELIGMANN ON DEATHS [ Feb. 19,
meshed cedematous tissue; so that probably cardiac failure is the
direct cause of death of many of these birds, the condition perhaps
being due to lack of exercise. One case of particular interest as
bearing out this view has occurred during the past year. While
some Zebras were being moved into the new enclosure in the
neighbourhood of the Seal pond,an Emu (Dromews nove-hollandie),
so placed as to be able to see what was happening, became
extremely excited, and running round its paddock either struck the
railings and collapsed against anes or else collapsed and fell against
the railings. In any case it could hardly have been injured by the
blow, since its feathers were not damaged or its skin torn. On
picking it up it was found to be quite dead, but at the post-mortem
examination no sign of disease could be found in the brain,
or abdominal organs, nor was the heart notably flabby.
Arterial Disease-—The aorta showing many aneurysms of an
old Tigress, which had lived in the Gardens for 1323 years, was
shown at one of the evening meetings of the Society, and a brief
note upon the condition appeared in the ‘ Proceedings’ *. An
illustration has since been prepared which is reproduced in text-
fig. 61, and shows the unusual condition of the vessel, the previous
description of which may be quoted here :—‘ The aorta shows
advanced arterial disease, most pronounced in the descending
aorta, where there is marked atheroma and where, in a length of
about 180 mm., there are 14 aneurysmal swellings varying in size
from that of a pea to that of a fair-sized plum. The two largest
swellings, the walls of which are of stony hardness, occur close
together on opposite sides of the artery.” Arterial disease, though
not common in the animals in confinement in the Gardens, cannot
be said to be rare, whereas in wild animals this disease is generally
considered to be very.rare. Perhaps an interesting parallel may
be drawn in this respect between man under civilised conditions
and animals in confinement. In the former arterial disease is of
course common, but there is considerable evidence against its
occurrence among certain people but just emerging from the Stone
Age.
Gastric Ulcer.—The experience of this and last year seems to
show that if gastric ulcers are not entirely confined to the
Carnivora and Marsupialia, they are at least most common in
these orders, since of a total of nine cases four occurred in
Carnivora and five in Marsupialia. A young Ocelot (felis par-
dalis) presented the lesions of this disease in a_ particularly
interesting form. On opening the belly, part of the colon looked
darker than usual, and it was found that this and the rectum
were full of dark semi-digested blood; there were many small
recent hemorrhages in the stomach, over some of these the mucosa
was destroyed but the ulcers had not penetrated deeply. In the
duodenum there were, however, sixteen orifices, all more or less
circular, and varying In size from that of the head of a large pin
to that of a threepenny-bit. One of the largest of these extended
* P.Z.S. 1906, p. 634.
Ix
1907. ] IN 'THE SOCIETY'S MENAGERIE. 173
as deeply as the serous coat, the others were less deep and
involved only the mucosa and a part of the muscular walls of the gut.
The Marsupials with gastric ulcer were all herbivorous.
Perforation of the Uterus.—An instance of this unusual accident
oceurred in a bitch of the North-African Jackal (Canis anthus).
The mammary glands betokened recent activity, and on opening
the abdomen the uterus projected well above the pelvis; there was
general peritonitis and the uterus itself was dark and intensely
congested. Within it lay a dead and extremely offensive fcetus,
while a small circular perforation existed in the uterine wall just
above the cervix on the left side, and it was apparently due to
leakage through this hole that peritonitis had set in.
New Growths.—The rarity of new growths referred to in last
year’s note is confirmed by this year’s post-mortem examinations
on a larger number of animals. Only four cases occurred ; two of
these, both in mammals, were carcinomata and presented no
unusual features; but the third case, a Bear, had multiple small
angeiomata of the liver, none bigger than a sixpenny-bit, which
cannot have exerted any evil effect on the health of the animal.
The last case occurred in a Chilian Pintail (Dafila spinicauda)
alleged to have been bred in the menagerie and to be 26 years old.
On opening the bird a mass the size of a turkey’s egg was seen
occupying the right flank in the front of the belly; it was not
adherent to the intestine or other organs but was enclosed within
a thin, loose capsule resembling peritoneum within which, with
the mass, were the supra-renals, which were not affected. The
mass could not be traced to any organ, but seemed to arise at the root
of the mesentery; no testes could be found, although the trachea
was of the normal male type; there was a white mass about the
size of a filbert in the right lobe of the liver, and both kidneys
were whitish and much enlarged. Microscopically it was seen that
the main mass was structurally a carcinoma, as was that in the liver,
and both these masses resembled the much enlarged kidneys which
were themselves carcinomatous.
My thanks are due to Mr. 8. G. Shattock for the diagnosis
of this remarkable case, which it seems must be classed as an
example of diffuse carcinomatous growth in both kidneys with
secondary masses in the liver and glands at the root of the
mesentery.
While considering the occurrence of new growths in birds,
allusion may be made to two very interesting conditions which
have recently come under my notice. The subject of the first of
these was a domestic fowl belonging to Dr. R. N. Salaman *
which died suddenly. On opening the belly cavity this was found
to be full of blood, and the liver was found to be diffusely angeio-
matous. The second condition occurred in a Sparrow examined by
Dr. A. Wilson * in the course of certain physiological work. On
opening the skull a spherical mass about the size of a pea was seen
* My thanks are due also to these gentlemen for allowing me to refer to their
specimens.
174 MR. J. T. CUNNINGHAM ON [Feb. 19
to press upon the right half of the cerebellum, which it had to
some extent excavated. This mass was enclosed in a sheath of
pia mater continuous with that over the posterior part of the
right cerebral hemisphere. Dr. Wilson thought that the spherical
mass might at one time have been joined to the cerebral hemi-
sphere, but of this there was no direct evidence. Microscopically
the tumour mass contains true nerve-cells, and so is an example of
an extremely rare condition.
3. On a peculiarly Abnormal Specimen of Turbot.
By J. T. Cunnincuam, M.A., F.Z:8.
[Received January 23, 1907. |
(Plate XI.*)
The specimen here described was sent tome by Dr. EH. J. Allen,
Director of the Marine Biological Laboratory, Plymouth, in the
beginning of December 1906, with a request that I should examine
and describe it. With the fish was a normal specimen and two
letters referring to them—one from Mr. John D, Enys, the other
from Miss Olivia L. Fox. Mr. Enys’ letter is dated Nov. 3, 1906,
and states that Miss Fox had then alive ina glass globe two
small Turbot caught on the sands at Polzeth, near the Doom Bar
at Padstow, on the north coast of Cornwall; that the abnormal
fish was dark on the under side and white on the upper. Miss Fox’s
letter states that she had had the fish about a month, and that the
upper side “‘ was becoming pigmented” since she first obtained it.
The specimen is 4:4 em. in length and presents a condition
which, so far as I am aware, has never previously been observed
or described in flat-fish of any species. With respect to the
position of the eyes, the fish is a reversed specimen—that is to
say, both eyes are on the right side, whereas normally in Turbot
they are on the left. With respect to colour, on the contrary
the specimen partially resembles a normal Turbot. The right side
is almost entirely unpigmented ; the greater part of the left side
is coloured like a normal Turbot. The pigmentation does not
extend uniformly over the whole of the left side, but is absent
from the head, and from the anterior part of the dorsal region
above the head. On these areas there are only a few scattered
black chromatophores. On the right or uncoloured side there are
also scattered black chromatophores rather more numerous than
on the left side of the head. It is important to note that the head
and anterior region of the right side, although not fully pigmented,
have more pigment than the rest of that side; between the eyes
and around the dorsal eye pigmentation is almost complete.
The number of dorsal fin-rays in the specimen is 65, of the
ventral 47. The characteristic tubercles of the adult Turbot are
not yet developed, but there are three little projections at the
base of each of the dorsal and ventral fin-rays, and also projections
* For explanation of the Plate, see p. 181.
Ie eo Wi. Ih. 2:
d.Greeen hth.
J.T. C del.
ABNORMAL YOUNG TURBOT.
1907.] AN ABNORMAL TURBOT. 175
at the bases of the caudal and ventral rays. These are probably
the beginnings of marginal tubercles.
The anterior end of the dorsal fin and the basal tissue which
carries it forma projecting hook-like process over the dorsal eye—
that is, the originally left eye, which has moved to the right side
of the head. This projection, due to the absence of attachment
between the base of the fin at the anterior end and the head,
occurs commonly in ambicolorate specimens of the turbot and less
frequently in ambicolorate specimens of other species of Plewro-
nectide (see Cunningham and MacMunn, “ Coloration of Skins of
Fishes, &c.,” Phil. Trans. 1894),
A letter from Miss Fox to Dr. Allen, dated Jan. 7, 1907,
explains that the fish was caught on Sept. 28 last year, and
lived in captivity till Nov. 28, when it died from some unknown
cause. When caught it was, unlike all the others seen at the same
time, quite stationary on the sand, which Miss Fox thought might
imply a certain blindness. In captivity, however, it was very
active, and certainly saw food very quickly, so that there is no
reason to think the function of the eyes was affected.
In Plate XI. I have figured the two sides of the abnormal
specimen. The normal specimen was 4°2 cm. long. Its meta-
morphosis is complete, but there are still a few scattered black
chromatophores on the right or lower side. Similar black
chromatophores are present on the right side of the abnormal
specimen, and they appear to be larger and slightly more
numerous ; but the difference is slight, so that the exposure of
the right side to light during the two months it was in captivity
had not produced much effect.
It seems to me that the only way to attempt an explanation of
the condition of this specimen is to base the explanation on the
view of the constitution of the ovum which was developed by
Weismann, and which is adopted in the Mendelian doctrine of
heredity. If the right side of the anterior or cephalic region were
more completely pigmented, we might regard the fish as consisting
of an anterior smaller part which was reversed, and a posterior
part which was normal. The condition would then be explained
by supposing the specimen developed from an ovum consisting of
parts usually occurring in separate ova. We know that reversed
specimens occur in various species of flat-fishes, e. g. the Flounder
(Pleuronectes flesus). In this species in some localities reversed
specimens are not only common, but abundant. At Plymouth I
found about 30 per cent. of the specimens captured had the eyes
and pigment on the left side instead of on the right. It is neces-
sary here to consider the precise terms to be used to indicate the
structural peculiarities which present themselves. It has been
usual to speak of a Flounder with eyes on the left side as reversed ;
but if we use the substantive corresponding to this adjective,
namely reversion, we are using a term which has been employed
in an entirely different sense, namely as meaning atavism, or the
recurrence in a species of some more or less remote ancestral form.
176 MR. J. T. CUNNINGHAM ON | Feb. 19,
It is true we might use the word reversal, but this is not suffi-
ciently distinct. In order to avoid confusion it will be better to
coin a new term, and it seems to me the most appropriate term is
“‘metastrophe,” meaning a change in the direction of the turning.
For adjectives we may use merely sinistral or dextral, referring to
left or right side, or for the abnormal condition in general we may
use metastrophiec.
Since, then, metastrophe frequently occurs in flat-fishes, and is
a congenital abnormality due to some abnormality in the constitu-
tion of the ovum, it is intelligible that it should occur in one part
of a fish and not in another. We may suppose the abnormality
in the whole fish is due to the interchange of position in the ovum
of the parts corresponding to the left and right sides of the body.
The abnormality does not, however, affect the viscera, which, as I
have pointed out in the memoir already cited, are constant in
position whether the fish is dextral or sinistral. In the particular
specimen of Turbot which we are considering, the head is dextral,
or metastrophic, the posterior portion normally sinistral, and its
origin is to be attributed to a corresponding abnormality in the
constitution of the ovum from which the fish was developed.
With regard to the question of the origin of such abnormalities
in the ovum, they may arise either in the cell-divisions which
occur in the multiplication of ova or spermatozoa of gametes,
to use the general term, or in the process of fertilisation, the
conjugation of the gametes. It might be suggested in this
particular case that the condition was due to a “‘ cross” between
an abnormal dextral specimen and a normal sinistral specimen,
the condition of the head-region being inherited from one parent
and that of the posterior region from another. But metastrophic
or dextral specimens are, so far as my experience goes, rare in the
Turbot, and it seems equally possible that the peculiar condition
of the gamete which gave rise to the abnormality was not due to
the condition of one of the parents.
Tt is not necessary to suppose that both of the gametes which
produced the fertilised ovum were abnormal : abnor mality in one
only may have been sufficient to produce the abnormality of
development. In the division of the gametes within the repro-
ductive organ of a parent fish, the chromosomes of the nucleus,
which are supposed to be the “carriers of heredity ” or to contain
the “determinants” which produce the characters of the organism
to which the gamete gives rise, normally divide severally so
that two similar ova are produced. In the final or reduction
division each chromosome does not divide, but the group of
chromosomes separates into two groups. In one or other of
these divisions the determinants might be displaced, so that either
all or some of those belonging to the left side were on the right
and vice versd, and thus a metastrophic gamete would be produced.
One important question that arises from the condition observed
in the specimen under discussion is, what bearing it has on the
experiments carried out by me some years ago at the Plymouth
1907. ] AN ABNORMAL TURBOY. Ti
Laboratory, and described in the paper already cited. In those
experiments pigment was developed on the lower sides of Flounders
as a result of the incidence of light. Here we have a specimen
of Turbot in which the upper side is exposed to light and is not
pigmented, while the lower side is pigmented. But it must be
noted that no adult specimen has been observed in which this
condition occurs. According to Miss Fox’s letter quoted above,
the upper side of this young turbot had already acquired some
pigment during the two months in which it lived in her possession.
It is quite possible therefore that if the specimen had lived to
become adult, the upper or right side would have become fully
pigmented in consequence of the action of light, and then the
specimen would have been exactly similar to other ambicolorate
specimens of Turbot, except that it was metastrophic, the eyes
being on the right side instead of the left.
In my experiments, I showed that when young fish in process
of metamorphosis were placed in the apparatus so that light fell
on the lower side and not on the upper, the normal hereditary
changes were not arrested, pigment disappeared from the lower
side as under normal conditions, and it was only later, after
long exposure to light, that pigment was developed on the lower
side. Thus, as the specimen we are here considering had not long
passed its metamorphosis, there is nothing inconsistent with my
results in the absence of pigment from the tight side, although that
side is uppermost and had been exposed to light for a short time.
The condition of the specimen here described suggests that the
usual ambicolorate abnormality is due also to partial metastrophe,
but that in these cases the anterior part of the body is normal or
sinistral, and the posterior part dextral. This view would explain
the remarkable fact, of which hitherto no explanation has been
given, that in the great majority of ambicolorate Turbot the lower
or right side of the head is unpigmented, just as in the specimen
here described the left side of the head is unpigmented. The
limits of the pigmentation are not absolutely constant. In the
majority of specimens which I have seen, the pigmentation
extends on to the lower jaw and the anterior end of the dorsal fin,
while the rest of the head in front of the preopercular bone is
unpigmented. One specimen in my list, however, had pigmenta-
tion over the whole of the lower side, including the head. If the
explanation suggested is correct, it follows that the young of an
ambicolorate specimen immediately after metamorphosis is without
pigment on the postcephalic portion of the upper cr left side, and
that it becomes ambicolorate in adult life in consequence of the
development of pigment on that side under the influence of light.
There is at present no direct evidence of this beyond the occurrence
of the specimen described in this paper, and the question must be
further investigated by the examination of large numbers of
young specimens. When pigmentation extends over the whole of
the lower side, mcluding the head, it cannot be said that the head
of the fish is normally asymmetrical; therefore the theory of
Proc. Zoou. Soc.—1907, No. XII. 12
178 MR. J. 1. CUNNINGHAM ON [Feb. 19,
partial metastrophe does not apply. In this case we must conclude
that some other explanation is to be sought, or we may suppose
that the boundaries between the determinant groups in the ovum
are not definite, and that the pigment determinants displaced to
the right side have extended to the head-region.
It may be objected that the persistence of colour on the lower
side of an ambicolorate Turbot is inconsistent with my views of
the action of light, that if pigment were produced on the upper
side it ought to disappear from the side turned to the ground.
This objection is of little weight, for my experiments show that it
is easier by means of light to produce some pigment where it was
previously absent, than to abolish it when it is present, by cutting
off the light. This is what might be expected, for in the evolution
of a flat-fish pigment has only recently disappeared from the lower
side, in consequence, as I believe, of the absence of light; and
therefore the pigmentless condition is not very strongly inherited,
and pigment is produced after a comparatively short exposure to
light. The positive character on the other hand, the presence
of pigment, has existed not only since the flat-fish was evolved,
but in a long line of ancestors before that, and therefore it would
probably take several generations to cause the pigment to disappear
completely by cutting off the light. It is quite possible that when
the lower side is congenitally pigmented, some proportion of the
pigment is lost in consequence of the absence of light, but such a
loss would not be obvious to observation and would be difficult to
demonstrate. Obviously a small amount of pigment appearing
where there was none before 1s evident at once, but the disappear-
ance of a small proportion from a strongly pigmented surface
makes no apparent difference to the colour, and there is no means
of measuring the amount of pigment for comparison in different
cases. There can be no doubt concerning the presence of a single
sheep in a field, but 1t is much more difficult to decide whether
there are a thousand or 999 in a flock.
Tt has long been known that in Pleawronectide generally, and
especially in Rhombus maximus, there is a marked correlation
between ambicoloration and the malformation of the dorsal fin
which occurs in the specimen described in this paper. It seems to
be generally supposed that in such specimens the dislocated eye
has not completed its change of position, and being on the edge
of the head instead of on the upper side, prevents the usual growth
forwards of the base of the dorsal fin. The condition is regarded
then as, like the ambicoloration, a reversion on the part of the
eyes and skull towards the primitive symmetry. Although I have
not fully investigated the structure anatomically, it 1s my opinion,
from external observation, that the eyes and skull are normal and
that the peculiarity is merely due to a want of that attachment
which normally occurs between the base of the fin and the skuil,
along the united ectethmoid or prefrontal, and frontal bones. T he
view L have suggested seems to me to give a better explanation
of this abnormality than has hitherto been proposed. If the head
1907. ] AN ABNORMAL TURBOT, 179
is metastrophic and the poster lor region normal, as in the specimen
here described, or vice versd, aS in ambicolorate specimens pre-
viously described, then the normal relation of the determinants of
these parts in the ovum, and therefore in development, is wanting.
The anterior end of the dorsal fin belongs to the posterior of the
two portions abnormally joined in the fish. It tends to grow
forward, but in the normal case in doing so unites with the right
side of the skull (in the Turbot); whereas in the abnormal specimen
here described, where the head is metastrophic, it has the left side
of the skull opposite to it, and with this side it has no congenital
relations, and so remains separate from it. In the more usual
case, where the eyes are on the left side as usual but the fish is
ambicolorate, a similar explanation would apply. Here the right
side of the skull is opposite the fin, as in the normal fish; but the
fin being itself metastrophic, the normal relations between fin
and skull in development are disturbed, and consequently they
remain separate. It may be said in fact that in all these cases
the fish. or the ovum from which it develops, is composed of two
separate parts united in an abnormal relation to one another in a
plane transverse to the long axis of the fish. Consequently the
normal continuity between the head and body is, as it were,
imperfect ; and in all probability this is the real reason why in
these cases the anterior end of the dorsal fin remains unattached
to the head.
The abnormality of the dorsal fin does not occur in specimens
which are entirely metastrophic. Here, although the characters
of the right side develop on the left, and vice versa—that is to
say, the determinants of the right and left sides have changed
places—the dislocation of determinants in the gamete has taken
place along the median plane, and therefore the longitudinal
continuity between fin and skull is not disturbed.
It is important to mention that the abnormality of the fin in
the specimen here described is not merely due to incomplete
metamorphosis. The normal specimen of the same size, or rather
smaller, sent with the abnormal, and captured at the same time,
shows complete metamorphosis, and in it the dorsal fin extends
forward attached to the head to a point anterior to the eyes.
The correlation between ambicoloration and the abnormality of
the dorsal fin isnot invariable. Cases occur in which ambicolorate
specimens are in this respect structurally normal. In the Phil.
Trans. memoir by myself and Dr. MacMunn (referred to above,
p- 175), I made the generalisation from the specimens of Turbot
then known to me, that if pigment was present over the whole of
the body behind the pre-opercular bone, and also on the lower jaw
and the anterior end of the dorsal fin, the malformation of the dorsal
fin was present; whereas if the pigment was less than this, the
malformation was absent. On the hypothesis of the cause which
T have suggested, the absence of the malformation inthe latter case
is intelligible, for then the junction between the metastrophic and
the normal parts of the body may be supposed to occur not between
12s
180 ON AN ABNORMAL TURBOY. [ Feb. 19,
the dorsal fin and the head, but in the course of the dorsal fin
itself. The anterior end of the fin then belonging to the same part
of the composite fish as the head, the relations of the two struc-
tures in development are not disturbed. Holt *, however, records
aspecimen of Turbot in which there was some pigment on the lower
side of the jaws and on other parts in front of the preoperculum,
and yet the head and fin were structurally normal. ‘This excep-
tion is not enough to disprove my hypothesis, for it may happen
exceptionally that the two parts in the composite gamete are so
exactly fitted together in their new relations, that the fin attaches
itself to the skull in development as in a normal specimen.
The fact which gives most support to my hypothesis, is that in
the great majority of ambicolorate Turbot the lower side of the
head is white and destitute of pigment. The same condition has
been seen by me in two specimens of Pleuronectes macrocephalus
and one specimen of Plaice, P. platessa. Inthe Flounder (P. flesus),
however, I have not found the absence of pigment from the lower
side of the head in ambicolorate specimens. In the Phil. Trans.
memoir I have recorded four specimens in which the lower side
was completely coloured and in which the usual abnormality of
the dorsal fin was present.
There is, however, another fact concerning ambicoloration
which is difficult to reconcile with the view that it is due to the
metastrophe of the posterior region, namely, that in ambicolorate
specimens not merely both sides are pigmented, but both sides are
equally armed—that is to say, the scales, spines, or tubercles are
equally developed on both sides. In normal flat-fishes the arma-
ture is much reduced on the lower side. In the Turbot the
scattered tubercles present on the upper side are almost entirely
absent from the lower side ; in the Flounder the rough spiny scales
along the lateral line and along the bases of the dorsal and ventral
fins are absent from the lower side. In ambicolorate specimens
the armature is not only present on the lower side, but also on
the upper; the abnormal specimens are not merely ambicolorate,
but ambiarmate. If, according to the hypothesis I have suggested,
the postcephalic region were metastrophic, the upper side should
be originally not merely without pigment, but without armature,
like the lower side in normal specimens. If the pigment on the
upper side in ambicolorate specimens were due to light, the arma-
ture should remain absent, unless the action of light produces
armature as well as pigment, for which there seems no reason and
for which we have no evidence. Something might be attributed
to the absence of friction from the upper side, but this is excluded
by the strong development of the armature on the lower side in
these abnormal specimens. These considerations are in favour
of the alternative hypothesis to explain ambicoloration and ambi-
armature, namely, that in the gamete two right or two left sides
are united instead of aright and left. In this case of course all
* E. W. L. Holt, “Studies in Teleostean Morphology from the Marine Labora-
tory at Cleethorpes,” P. Z. S. 1894, p. 413.
1907. ] ON THE AZYGOS VEINS IN MAMMALS. 181
the characters of the upper side would be reproduced in the lower,
both pigmentation and armature. The symmetry does not extend
to the eyes and skull, but then the malformation of the head may
be due to a partial symmetry, a reduction of the normal asymmetry.
It is possible that both cases occur in different specimens—that
is to say, that in some ambicolorate specimens the condition is
due to metastrophe of the posterior region of the body ; in others
to secondary symmetry, the doubling of the upper side in the
gamete. This cannot be decided without further investigation of
abnormal specimens both in the young state and theadult. There
can be no doubt of the importance of the unique condition
exhibited by the specimen here described, or that its condition
is best explained on the view I have suggested, namely, that it
consists of a metastrophic head joined to a normal body.
I beg to offer my thanks and congratulations to Miss Fox and
Mr. Enys for making the specimen known, and my thanks to
Dr. Allen for allowing me to describe it.
EXPLANATION OF PLATE XI.
Abnormal Young Turbot.
ea)
39
.1. Right (upper) side of abnormal Turbot 4:4 mm. long, enlarged. The eyes
are on the right side of the head, and the dorsal fin projects anteriorly as
a free process. Some pigment on the head dorsally, elsewhere only
scattered black chromatophores.
g. 2. Left (lower) side of the abnormal Turbot shown in fig. 1. No eyes on
the lett side of head, nor pigment. Pigment on the posterior region as in
normal specimen.
=
8
4. On the Azygos Veins in the Mammalia. By Frank
BE. Brpparp, M.A. (Oxon.), F.R.S., Prosector to the
Society.
[Received February 1, 1907. |
(Text-figures 62-73.)
CONTENTS.
(1) Introductory, p. 181. (9) The Azygos and other thoracic veins
(2) The Azygos Veins in the Ungulata, | i NG vous ae UI gojpouciems
p. 183 coypu, p- 213.
eee ee 196 (10) The condition of the Azygos and
(3) Insectivora, p. ; | the Classification of Mammals,
(4) Lemurs and Apes, p. 197. p. 218.
(5) Edentata, p. 198. | (11) The position of the Azygos with
pel RO A | reference to the Vertebre, p. 219.
a oes a ne (12) The Ven Intercostales Supreme,
ar als, p. 2UU. 92
2 p- 221.
(8) Rodents, p. 208. (13) Conclusions, p. 222.
(1) Introductory.
It appeared to Cuvier*—and presumably to his editor Duvernoy,
since the statement is left unannotated—that the Azygos veins in
mammals were too variable to offer zoological characters of value ;
* Anat. Comp. ed. 2, t. vi. (Paris, 1839) p. 238,
182 MR, F. E. BEDDARD ON THE | Feb. 19,
for he wrote, “ L’insertion de azygos, Vexistence d’une azygos du
coté gauche sont assez variables; mais on sait que les mémes
circonstances varient dans VPhomme. Elles ne méritent pas con-
séquemment de nous arréter.” Muilne-Edwards*, on the other
hand, and rightly, treats these veins as of importance, and tabu-
lated the main variations. These statements really express the
main facts as we know them today. With reference to the
Marsupials, indeed, the assertion of Milne-Edwards seems to me
to be nearer to the truth than some generalisations made more
recently. This group is placed by Milne-Edwards under the
heading “ Les deux veines azygos également développées.”
Although I shall show reasons for slightly criticising this state-
ment, it is not greatly exaggerated. On the other hand,
denying the existence of a right azygos (“ Point de veine azygos
«1 droite”) in the Sheep, Ox, Goat, Chevrotain, Pig, and Tapir he is,
in my opinion, not by any means so accurate. With regard to
the Tapir there is some error in Milne-Edwards’s statement ; for
he also places that animal under the heading of those animals
which only possess a right azygos. Sir Richard Owen’s classical
text-book, published ten years later than the volume of Milne-
Edwards's great work, adds but little to the record of facts con-
cerning the azygos veins. There are, however, numerous scattered
references to the condition of these veins in various mammals by
Owen and others, to some of which I am able to refer in the
course of the following pages.
Many of these papers are quoted by Hochstetter? in his
memoir dealing with the development of the Azygos (and other
veins) in the Mammalia. A large number, however, relate to
the condition of the azygos in man, and I do not attempt here to
follow up that very large subject. I limit myself to such other
mammals as I have been able to dissect, in many of which the
azygos has not been described. The classificatory importance of
the azygos has been recognised by Dr. Max Weber {, and there is
no doubt that its conditions are often distinctive of genera or of
whole orders of mammals. I propose, however, to deal with this
matter after exposing the facts which [ have gathered together
by degrees during several years of intermittent work upon the
subject, which is a larger collection of facts concerning this vein
than has previously been br ought together.
Opinion with regard to the morphological nature of the azygos
veins has lately undergone some change. Until lately the pre-
valent view was that one or both of the postcardinals persisted as
the Azygos, the Hemiazygos, or both. This view is embodied in
diagrams in many text-books.
Of recent writers Messrs. Parker and Tozier§ appear to hold
* Anat. et Phys. Comp. vol. iii. (Paris, 1858) pp. 595-598.
+ Morph. Jahrb. xx. 1893, p. 642 &c. Milne-Edwards’s résumé is largely based
upon the observations of Bardeleben (Arch. f. Anat. u. Phys. 1848) and Marshall
(“ Development of great Anterior Veins &c.,” Phil. Trans. 1850).
t Die Saugetiere, Jena, 1904.
§ “Postcardinal Veins in Swine,” Bull. Mus. Comp. Zool. vol. xxxi. 1898, p. 133.
1907. | AZYGOS VEINS IN MAMMALS. 183
much the same view. For they remark that in the Pig “the
hemiazygos” (which is the left azygos of my nomenclature) ‘‘ from
the region of the heart to the tenth rib is therefore to be
regar ded as the persistent anterior portion of the left post-
cardinal.” The rest of the vein is formed, as they think, from
“the accessory veins,” which appear to be the subcardinals of
McClure*. They thus agree with Rathke in holding that the
anterior part of the azygos is persistent posteardinal, but differ
from him as to the mode of formation of the posterior region of
the azygos; for Rathke held that this region was due to a con-
tinual longitudinal anastomosis between intercostal veins, and
was thus an entirely new structure. Zumstein tT put forward the
older view also in the case of man, where that anatomist believed
that he had traced the azygos and the hemiazygos to the post-
cardinals exclusively. In the Guinea-pig, however, he = found
that the posteardinals took practically no share in the formation
of the azygos. Hochstetter $ came to conclusions which were not
dissimilar. He allowed in the case of the Rabbit and of the Cat
that the azygos of the adult down to about the eighth thoracic
segment was the posteardinal, but that thereafter it was a new
structure not formed from the postcardinal vems: “von da an
caudalwirts aber ist sie eine Neubildung.” This region of the
posteardinal, in fact, becomes a part of the posteaval. a pertectly
different origin of the azygos veins is asserted by McClure || of
Didelphys. ‘Excepting just at their entry into the duct of Cuvier
they are quite independent of the postcardinals and of the sub-
cardinals, though connected with the former by cross anastomoses.
T shall bring forward various facts in the following pages which
bear upon this question of the morphological nature of the azygos
veins in mammals.
(2) The Azygos Veins in the UNGULATA.
I have paid special attention to this group since I have par-
ticularly favourable opportunities, as compared with those enjoyed
by other zoologists, of examining recently dead specimens. ‘These
bulky animals obviously cannot be preserved, and must therefore
be studied immediately after death. It thus follows that I am
able to add a good deal to what is known upon the subject.
It will be seen, however, that there is a very general agreement
among the Artiodactyle division of that Order as contrasting
with ‘the Perissodactyles, but that the division, as shown by
the azygos vein, is not absolute. Max Weber, in lis recent text-
book (Die Siugetiere, p. 642), uses the condition of the azygos
to define the Artiodactyla thus :—‘ Die Vena azygos fehlt; die
* “ Development of Veins of Didelphys,” Amer. Jour. Anat. v. 1906, p. 163.
ii “ Bntwicklunge des Venensystems des Menschen,” Anat.-Hefte, Bd. vi. 1896.
+ “ Venensystem bei dem Meerschweinchen,” ibid. Bd. vin. 1897.
§ Loe. cit. p. 574. || Loe. cit. p. 185.
184 MR. F. E. BEDDARD ON THE [Feb. 19,
Vena hemiazygos miindet direkt oder indirekt in die Vorkammer.”
But, as I shall show, this is not quite universal.
Since the system of azygos veins is as perfect in the Gnu,
Connochetes g grit, as in any other Ungulata, and more elaborately
developed than in many, I shall commence with a description of
those veins in this Antelope. The anterior vena cava receives a
right and a left azygos vein which enter it very nearly, if not
exactly, opposite to each other. Of these two the right is rather
longer than the left. This right azygos receives five branches, of
which the most anterior is composed of affluents from two ribs.
It is evident therefore that the right azygos does not reach back
nearly as far as the diaphragm. The left vein is composed of
only four intercostal branches. The blood from the ribs lying
behind these four ate connected, however, into another lon-
gitudinal trunk lying on the left side. Seven or eight of these
branches coming from as many intercostal spaces combine to
form a vein running in the same straight line as the left azygos,
but not joining it anteriorly. The vein, in fact, opens inde-
pendently into the right auricle, as previous observers have noted
for other Ungulates.
It is not a little remarkable that the other species of Gnu,
viz. Connochetes taurinus, which I have also dissected, shows
differences in respect of these veins from Connochetes gnu. On
the right side the azygos is a trifle smaller and collects blood only
from four intercostal spaces. On the left side the azygos is very
much longer, with seven or eight affluents. It enters the vena
cava superior as in Marsupials &e., which possess two azygos veins.
I did not detect any branch putting this vein into direct com-
munication with the right auricle, such as occurs in Connochetes
gnu and in some other forms which I shall deal with immediately.
An intermediate condition is offered by another Antelope, Bubalis
caama. In this animal the right azygos is composed of five
affluents. On the left side we have both the anterior and the
posterior vein of Connochetes gnu, and, as in that animal, the
latter communication with the auricle direct. But a slender twig
connects the two veins, which are thus a continuous vessel as in
Connochetes taurinus.
The azygos veins of Rhaphicerus melanotis present the usual
Antilopine characters; but there are differences of detail from
those of some other forms. On the right side, the internal
mammary vein is followed in passing towards the heart by a
superior intercostal which divides into two branches, each of which
supplies the first or the second intercostal space. Behind this
arises the right azygos, which is composed of three afiluents ; the
first lies in the intercostal space 2/3, the others in 3/4 and 4/5.
Thus the intercostal space 2/3 has two veins. On the left side
there is a superior intercostal opening exactly opposite to the right-
hand vein; but it isa larger vessel though it only draws blood from
two intercostal spaces. The left azygos, as 1m alhed forms, 1s well
developed and enters the heart in common with the vena cava
1907. | AZYGOS VEINS IN MAMMALS. 185
inferior. It does not, however, consist only of a region lying
behind its point of entrance into the auricle ; but a branch also
runs forward supplying four intercostal spaces, the first being that
between ribs 2/3. The vein does not join the superior intercostal.
But it will be noted that rib interspace 2/3 has on the right two
veins.
Ourebia nigricaudata is much like Rhaphicerus. The vena cava.
anterior receives a pair of veins, the superior intercostals, which
bring back blood from the region of the first rib only. The two
veins are not absolutely symmetrical as to their point of entrance
into the vena cava. The second to the fifth rib inclusive supply
four intercostals which unite to form the right azygos. On the
left side of the body the azygos consists of an anterior and a
posterior section which unite at their communication with the
right auricle. The anterior vein is very slender, excepting in the
region at and near to its fusion with the posterior vein. The
azygos proper, 7.e. the posterior section, is a stout well-developed
vein which receives blood from the intercostals on both sides of
the body, It ends at the diaphragm in a bifurcation formed by
the two intercostals supplying the last (the thirteenth) ribs.
Cephalophus grimmii showed an identity of arrangement in two
examples, one a male, the other a female. The better developed
left azygos entered the heart in company with the vena cava
inferior, and not by a separate orifice. On the right side of the
body the azygos consisted certainly of four intercostal affluents.
The last two of these (at any rate in the male example) encircled
the fourth rib. C. maawelli shows much the same relation to
C. grimmii that the two species of Gnushow. For the left azygos
opens into the precaval. But it gives off a branch to the auricle.
Oryx leucoryx has also a comparatively short right azygos
composed, however, certainly of five intercostal branches. The
longer left azygos enters the heart either directly or with the
vena cava inferior. Above the influx of the right azygos is a
single inferior intercostal on the right side of the body, and on
the left side a corresponding vein which, however, immediately
divides into two trunks. There was, however, no connection
between the posterior of these two veins and the lower section
of the azygos system.
The arrangement of these veins is much the same in Oryx beatria
as it isin Oryx leucoryx. The left and principal azygos commences
with the interspace between ribs 6 and 5. Above this is a vena
suprema intercostalis, composed of only two branches lying in the
first two rib interspaces. To this corresponds exactly on the right
side a vein which draws blood only from the first intercostal space.
Behind this the right azygos opens into the precaval opposite to
the second or third rib. Its branches extend back to the sixth
rib, and they begin with the second intercostal space.
I have selected for figuring here (text-fig. 62) the azygos and
immediately related veins of Cervicapra bohor, on account of their
splendid condition in the male example which U had the opportunity
186 MR. F. E. BEDDARD ON THE | Feb. 19,
of dissecting in the month of December of last year. I am pretty
certain that I have been able to note down the characters of all the
veins of this system in the present species. The animal, I may
remark, was young and very slightly diseased. Cervicapra bohor
agrees in essentials with other Antelopes; but there are differences
Text-fig. 62.
Pp,
L
“fe-U.70
Scl.m Vg ace
Wie
IL /2 A
ae
2
:
=
SARE.
a
5)
Azygos veins of Cervicapra bohor.
Az.l. Left azygos; Az.r. Right azygos; 7.m. Internal mammary; Sc/Z. Subclavian ;
S.i. Superior intercostal ; p.c. Precaval. First three ribs numbered 1, 2, 3.
of detail which are worth recording. The main azygos trunk lies
as usual on the left side and opens into the heart. The level at
which it bends towards the heart is opposite to the seventh rib.
1907.4 AZYGOS VEINS IN MAMMALS. 187
Its affluents, however, commence on the posterior side of the fourth
rib. Beyond the last pair of affluents, which lie behind the
thirteenth (and last) rib, the trunk is continued back as a very
slender vessel. The first intercostal artery which crosses this
azygos lies behind the tenth rib. Anteriorly, the second and
third intercostal spaces give rise to vessels which unite into a
trunk which debouches into the left subclavian vein. The right
Text-fig. 63.
Sel. pe. r7r7e-
= dj i) See.
eas ae
= He : oe
Aer e
; | ueere Az.t
oN
Azygos veins ot Gazella euchore. Lettering as in text-fig. 62.
subclavian, it may be remarked, joins the precava quite a con-
siderable distance behind the opening of the left subclavian—a
marked asymmetry which is not common. It receives, however,
no branches from intercostal spaces and enters the thoracic cavity
in the usual position—that is, in front of the first rib. Theright
azygos is of fair size; it enters the precava about opposite the
fourth rib. It receives branches from intercostal spaces 2 to 6
(inclusive).
188 MR. F. E, BEDDARD ON THE [Feb. 19,
In Gazella euchore (the example dissected was a male) the dis-
position of the several veins of the azygos system was as follows
and as is shown in the accompanying illustration (text-fig. 63).
The illustration in question shows that the subclavian veins are
symmetrical with each other and occupy the usual position,
emerging as they do in front of the first rib. To the left sub-
clavian is attached the corresponding vena intercostals supreméa.
This vein collects blocd from the second and third intercostal
spaces. I am not absolutely certain whether its longitudinal
trunk is not continuous with the left azygos. The latter com-
mences with a branch from behind the fifth rib; it enters the
heart on a level with the sixth or seventh rib. The right
azygos 1s fairly well developed and enters the precaval vein on a
level with the second rib. It collects blood by branches from the
second to the fifth intercostal spaces inclusive.
Tn a female Capra megaceros the left azygos also opened directly
into the auricle; its first branch arose from behind the sixth rib.
Anteriorly, a superior intercostal consisted of two branches
lying respectively between ribs 1 and 2 and 2 and 3. On the
opposite side a vessel corresponded exactly to this in point of
entrance into the precava, but drew blood only from the first
intercostal space. Immediately behind this opens the right LEA EOS,
which is made up of four intercostal branches from ribs 2 to 5.
In another example, also a female, the azygos was also on both
sides, but the right vein was smaller than in the specimen just
described and only drew blood from two intercostal spaces.
Hemitragus jemlaica ( ¢ juv.) is rather different in the arrange-
ment of these various veins. There is, on the left side, the typical
Artiodactyl azygos entering the heart directly and commencing
with a branch in intercosta i space 4/5. There is no superior inter-
costal on this side; and on the right side there is only a single
vein and that corresponds to the superior intercostal, since it
draws blood only from behind the second rib.
Of Nemorhedus swetienhami I have dissected a single male
example. As usual, the left azygos was the predominant vein
concerned with the intercostal circulation. It receives branches
from the fifth rib onwards, and the main trunk was traced some
way into the lumbar region behind the last (7. e. the thirteenth)
rib. The vena cava anterior receives the right azygos and
corresponding vein exactly opposite to it on the left side. The
former receives blood from the first six ribs and ends entirely at
the level of the sixth rib. The corresponding vein of the left side
has naturally fewer branches and ends at the fourth rib. I did
not ascertain whether there was any connection between the
anterior and posterior series of intercostal veins on the left side of
the body.
In an old example of Phacocherus cethiopicus (text-fig. 64) the
azygos veins were rather different from those of the two species
just described. The general plan was the same: that is, there is
a long azygos vein on the left side. This vein started with an
USO] AZYGOS VEINS IN MAMMALS. 189
affluent from the intercostal space between ribs 5 and 6. As far
as the twelfth rib the vein lay outside the intercostal arteries,
which up to this pomt passed between the azygos vein and the
vertebral centra. The intercostal artery arising from the aorta
between the twelfth and thirteenth ribs passed to the outside of the
azygos. Asa rule among mammals this point, where the azygos
changes its position relatively to the intercostal arteries, is further
Text-fig. 64.
mB P.c.
Az.
Azygos veins of Phacocherus ethiopicus. Lettering as in text-fig. 62.
forward. Anteriorly the precaval vein receives two quite sym-
metrically disposed veins, one from each side of the body, which
convey blood from the more anterior intercostal spaces. ‘These
veins, moreover, are not only symmetrical with each other in so far
as regards their point of junction with the precava: they draw
blood from exactly the same intercostal spaces. Each consists of
190 MR. F, E. BEDDARD ON THE [Feb. 19,
four branches supplying the first four ribs. Their point of
opening is about opposite the second rib.
In a young example of the African Red River-Hog (Poéamo-
cherus africanus) the azygos was also limited to the left side of
the body, as in Porcula salvania. In Potamocherus, however,
this vein enters the heart as in Ungulates; while in Porcula it
seems to have the relations of the azygos in Marsupials &c., where
a left azygos 1s present. The difference is not really of great
importance as indicative of an anomaly in one or the other of these
two genera of Suidee; for we have in other Ungulates—e. g., in
Connochetes gnu—a left azygos which is divided into two parts
whereof one opens into the jugular and the other into the heart.
One or other arrangement is found in each of the Suine genera
Porcula and Potamocherus. Potamocherus shows the arrange-
ment which was found in Sus scrofa by Hunter * and quoted by
Owen in his ‘Comparative Anatomy’? in dealing with the
azygos veins of mammals.
In the Pygmy Hog (Porcula salvania) the azygos vein is only
developed upon the left side of the body ; there was absolutely no
trace of this vein that I could discover upon the right side. It is
large and thick, and its branches are important and obvious with
the exception of the first. This more slender branch divides on
issuing from the main trunk into three twigs, of which two run
on each side of a rib. After this there are seven branches, which
are large and were turgid with blood in the individual examined.
These branches correspond of course regularly to the ribs; but
they supply eight intercostal spaces, since the last of them, in
which the azygos ends, bifurcates over its rib. The azygos Ties
very definitely upon the left side of the aorta ; it is not median in
position as is so often the case with this vem. The vem ended
in front of the diaphragm ; nor could I detect any branch, however
thin, which continued on the main trunk behind the diaphragm in
the direction of lumbar veins or of the postcaval or renal veins.
These observations refer to one example only, which was a female.
The Musk Deer (dZoschus moschiferus) presents us with a system
of azygos veins like those of other Ungulates. On the left side
the well-developed azygos enters the aur ‘icle directly. On the right
side the small azygos consists of one vein only. In front of this,
and on both sides of the body, is a superior intercostal.
Dorcatherium aquaticum did not show, as might perhaps have
been expected, an arrangement of a specially primitive character.
It is much like other Ungulates. There is a small azygos
on the right side of the body formed by three aftluents only.
The azygos of the left side only communicates with the right
auricle. It is as extensive as is usual among Ungulates and con-
sists of many intercostal affluents. The first of these branches is
made up of two tributaries. There is no anteriorly running
* ©Hssays and Observations on Natural History &e.,’ arranged by Richard Owen,
vol. 11. 1861, p. 124.
+ Vol. iii. 1868, p. 555.
1907.| AZYGOS VEINS IN MAMMALS. 191
extension of this vein such as oceurs in, for example, Rhaphicerus.
It is interesting to note that Dorcatheriwm does not agree with
its nearest ally, Vragulus, as the following account of the latter
genus will show.
Of Tragulus meminna (text-fig. 65) I have been able to examine
two adults—one of each sex—and a young one, nearly ready for
Text-fig.65.
Ga Sc.
4 SLE = 7
iV | f | 2
bf ‘
ART pe
A
/f
——
py
vi
Hay
A
//
ead
| |
Post. C. [oS |
Azygos veins of Tragulus meminna.
post.c. Remains of posterior cardinal of right side (?).
Other letters as in text-fig. 62.
birth, taken out of the uterus of the female. In all three the
azygos was present only on the right side and entered the precaval
very near to the heart. The nearness to the heart seemed to me
192 MR. F. E, BEDDARD ON THE [ Feb. 19,
to be more marked in the two adults than in the feetus. In the
male adult the first affluent of the azygos arose behind the fourth
rib; after this there was a gap of one rib, and three very much
stouter affluents arose behind the sixth, seventh, and eighth ribs
respectively. Thereafter the psoas muscles concealed the inter-
costal veins at their entry into the azygos, but they appeared to
be regular. In the female Deer the same very stout intercostal
veins arose from the intercostal spaces 6-7, 7-8, 8-9 as in the male,
but I did not find the slender anterior intercostal which I have
described in the male; I should not like, however, to assert that
it was absent. It is very interesting to observe this constancy in
a vein which has been stigmatised as most inconstant. Moreover,
the foetus showed the same arrangement of these veins with a
slight difference. The first affluent of the azygos was very stout
and arose between ribs 3 and 4, being thus a,rib further forward ;
but after this the first affluent of the regular series arose in the
same way as in the two specimens of 7ragulus meminna Just
described, behind the sixth rib. In this feetus, moreover, a very
slender intercostal, which I did not find in either of the adults,
arose from behind the second rib and either joined the precaval
independently or just at the point of entrance of the azygos. In
this fcetus the first rib sent an intercostal vein to the subclavian
of the right side. On the left side none of the three specimens
had any properly developed azygos. But in the foetus and the
male adult a large cesophageal vein received intercostal affluents
from the second and third ribs on the left side and entered the
subclavian of that side. There was something to correspond in
the female; but Iam unable to give a proper description. In any
case, the presence of rather more veins 1m the fcetus and the absence
of at any rate much variation in the azygos system of the adult
Tragulus meminna *, ave noteworthy. It is possible indeed that
there was no variation at all.
There is a very considerable agreement between the azygos vein
of Tragulus and that of Cervus sika, but not, as will be shown pre-
sently, of Cervus aristotelis. Inafemale of Cervus sika the azygos
was developed only upon the right side of the body. There were no
traces of anything of the kind on the left side. The right azygos
enters the precaval rather forward—in fact, opposite to the second
orthird rib. The first affluent which joins the azygos runs parallel
* An interesting note upon certain points in the venous system of this Deerlet
(“The Posteava of an adult Indian Chevrotain, Tragulus meminna Krxieben,”
Anat. Anz. Bd. xxix. 1906, p. 375), by Prof. McClure, has come into my hands
through the kindness of the author during the writing of the present memoir. In
this note it is remarked that, contrary to what is found in most mammals, but
agreeing with the conditions observable in Dasypus, Elephas, and the Marsupials
(generally), the postrenal division of the postcava lies directly ventrally to the aorta,
instead of dorsally and to one side. Dr. McClure naturally wonders if this is cha-
racteristic of Tragulus meminna, or is abnormal. I looked into the matter carefully
in the three specimens upon which I have reported above, and find that Dr. McClure
has discovered a perfectly normal character of this primitive Ruminant. I may
furthermore point out that he figures (Joc. cit. fig. 2) the rigbt renal vein as double
and the left as being single as well as much longer. I found this also to be the case
in the adult raale and in the fcetal male, but not in the adult female.
1907]. AZYGOS VEINS IN MAMMALS. 193
with it for some little distance as a longitudinal trunk, and receives
branches from the second to the fifth mtercostal spaces inclusive.
The main trunk of the azygos commences with an affluent from
the sixth intercostal space. There is, it will be observed, an
obvious likeness here to certain Rodents also; for instance,
Celogenys*. I may also observe that the vein lettered post.c. in
text-fig. 65 also probably corresponds to this additional azygos, for
the nature of which see p. 213 below.
In Cervus aristotelis the azygos on the right side of the body
pours blood into the single precaval from five aftluents, which
collect. blood from the first five intercostal spaces. On the left
side there is a corresponding vein which, however, flows into the
precaval rather in front of the right azygos, between the second
and third ribs. This collects blood from the first and a few sub-
sequent intercostal spaces. I think that it is continuous with the
usual Artiodactyle left azygos, which enters the heart opposite to
the sixth rib and extends back to the diaphragm.
Of Perissodactyle Ungulates I have only examined Lquus
chapmani. The arrangement here was precisely as in the Horse.
The azygos was present upon the right side only. The first
affluent contributing to the vein arose from the interspace
between the fifth and sixth ribs, and the vessel appeared to end
posteriorly after the affluent from the thirteenth rib.
Hyrax capensis.—This animal shows affinities to the Ungulates
in the structure of the azygos veins combined with a good many
differences from such Ungulates as I have been able to examine.
It is noteworthy that this ‘‘Subungulate” agrees with the Horse
and Tragulus vather than with most Artiodactyles in the much
greater size of the right azygos. In view of other comparisons
that have been made between Hyrax and the Perissodactyle
Ungulates, this additional fact is of interest. In the Rhinoceros,
according to Owen *, the right azygos is the principal azygos. This
feature alone is, however, “obviously not enough to establish an
affinity with the Per issodactyle section of the Ungulata, for many
mammals possess only the right azygos, with or w ithout traces of
the left. It is the characters of the azygos on the left side which
indicate the Ungulate affinities of this animal. In one of two
examples of Hyraa capensis which I have dissected, the first two
costal interspaces were occupied by veins which united to form a
single trunk opening into the vena cava anterior. These represent
the superior intercostal vein of other mammals. On the right side
of the body the corresponding veins were present, but each opened
separately into the vena cava. The next two ribs on the left side,
2. e. Nos. 3and 4, were supplied by a vein each, the two veins lying
both of them in the same intercostal space, between ribs 3 and 4.
These united to form a slender trunk which passed backwards
* See p. 209.
+ “On the Anatomy of the Indian Rhinoceros,” Trans. Z.S. iv. p.46. The Tapir
also has been stated to possess a large right azygos.
Proc. Zoou. Soc.—1907, No. XIIT. 13
194 MR. F. E. BEDDARD ON THE [ Feb. 19,
and opened into the vena cava inferior (text-fig. 66) just before
the debouchment of the latter into the right auricle. This is
clearly the Artiodactyle condition of the vein in question, though
differing in detail from that of any particular Artiodactyle that
has been studied and described. For in that group there is either
Text-fig. 66.
Sct.
aeons) (EE ae ees:
RNG)
LT Tre
Az.r-}
Hyrax capensis; two different arrangements of azygos system.
post.cav. Postcaval. Other letters as in text-fig. 62.
an anterior and a posterior branch of the left azygos which unite
close to the opening into the auricle or vena cava posterior, or the
posterior branch alone is present. In Hyrax it is the anterior
branch alone which is present.
1907.] AZYGOS VEINS IN MAMMALS. 195
The corresponding ribs on the right side were supplied with
veins joining the long right azygos. The azygos system of Hyra«
has been partly described by Brandt* in a memoir upon the
general anatomy of Hyrax, who observes that the vena cava
posterior enters the right auricle ‘ nachdem sie die vorderen
Vene intercostales und die Vena azygos aufgenommen.” ‘The same
fact, according to Weber 7, has been also noted by George =; but
I have not had the opportunity of studying this paper §. Nothing
is said by Brandt concerning the right azygos. It is, I presume,
legitimate to compare the descending region of the left azygos in
Hyrax with the left anterior cava. In a second specimen I did
not find a left azygos opening into the vena cava inferior. But
JT am not inclined to deny its existence. The large right azygos
was equally well developed in this example.
In a third example of Hyrax capensis (a female) the azygos
system was entirely developed upon the right side of the body,
and differed considerably from that of the two individuals just
described. The main azygos stem (text-fig. 66) was traceable
to the lumbar region, where it communicated with the vena cava
inferior (not opposite to the renal vein) in common with a lumbar
vein. The vein undoubtedly belongs to the right side since it
lies on the trachea on the right side of that tube. Its mode of
termination anteriorly is, so far as my own experience goes, unusual
among mammals. Instead of entering the jugular some little
distance in front of the heart, as is elsewhere (so far as | have seen
in my own dissections) invariably the case with the right azygos
when present, it enters that vein so near to its entry into the
auricle that it may almost be said to enter the a
But there is, of course, no question as to a direct communication
with the coronary sinus, like the left azygos of the Cavicornia. In
front of this. vein, which debouches into the heart opposite to the
fifth rib, is a small vein which receives blood from the 2nd and
3rd intercostal spaces.
The disposition of the azygos veins in the Artiodactyle Ungu-
lates seems, therefore, to be fairly constant in the group, though
the left azygos does not quite invariably open directly into the
right auricle or into the vena cava posterior just before the open-
ing of the latter into the right auricle. This arrangement of the
left azygos has not been found in any other group of mammals
excepting only in the Mole, where it has been stated to be the
same as in the Artiodactyles. Possibly this fact may be con-
sidered as requiring confirmation. It is, however, interesting to
note that as an abnormality the left azygos in man may open
into the right auricle directly. I have not attempted to compare
* Mém. Ac. St. Pétersbourg, ser. 7, t. xiv. p. 65.
+ Die Saugetiere, Jena, 1904.
+ Bibl. Ecole Hautes-Htudes, Sec. Sc. Nat. xii. 1875.
§ In an earlier paper by George (Ann. Sci. Nat. (6) 1. 1874) a right azygos only is
described. : J
|| This is also the case with the Horse, according to Chauveau and Arloing. Ihave
also described the same arrangement in Zragulus.
lai
196 MR. F. E, BEDDARD ON THE | Feb. 19,
the variations in the human subject with the conditions which
are normal in other mammals, and it is possible that many
examples have been recorded of this particular abnormality in
man. I refer only to one instance, which I cull from the list of
literature given by Hochstetter in his memoir upon the develop-
ment of the veins in mammals*.
(3) LNsECTIVORA.
My notes upon this group are very few, but not without
interest. Milne-Edwards has observed + that the Mole possesses
both azygos veins, and that the left debouches into the right
Text-fig. 67.
ie
AC)
a a ae oes
= {=e fe
We ee
S—-
ESS
Le,
eves
AN
Ad
He
{
Azygos veins of Hrinaceus algirus. Lettering as in text-fig. 62.
auricle precisely as in certain Ungulates. This fact is not referred
to by Owen, who however states + that “the left vena azygos
communicates with the left precaval in the Hedgehog and many
* Gruber, “ Ueber einen Fall von Einmiindung der V. hemiazygos in das Atrium
dextrum cordis beim Menschen,” Arch. f. Anat. u. Phys. 1864, p. 729.
+ Anat. et Phys. Comp. 11. 1868, p. 595.
+ Comp. Anat. & Phys, Vertebrates, vol. ii. 1868, p. 553.
1907. | AZYGOS VEINS IN MAMMALS. 197
others [of the “ Lissencephala”], and is larger than the right.”
This does not exactly agree with what I have found in an example
of Hrinaceus algirus, 9. Two azygos veins were present (text-
fig. 67) and of equal length. Each opened into the precaval of its
own side at a point corresponding to the interval between the
second and third ribs. In the ease of the right azygos, the first
affluent arising between the second and third ribs and the second
arising between ribs 5 and 6 were particularly large. It is note-
worthy that posteriorly there was also an irregularity in the
branches of the azygos, one branch often serving two intercostal
spaces. The specially thick affluents were not noticeable on the
left: azygos.
I have also dissected one example of the Common Hedgehog
(Erinaceus ewropeus), which differs both from Owen’s account of
that species already referred to and from Hrinaceus algirus as
dissected by myself. In this Hedgehog the right azygos alone was
well developed, and commenced with an affluent from the first rib
space, opposite to which it entered the right precaval. On the
left side a very small azygos was present consisting, so far as I
could see, of only one branch arising from the first intercostal
space. It joined the left precaval at a point apparently exactly
opposite the right azygos.
(4) Lemurs and APES.
Of this group I have examined a considerable number of species,
and in all of them the right azygos as a distinct vein is present,
and alone present. I have found this to be the case in the
following, viz. Lemur catta (3 examples), L. mongoz, L. xantho-
mystax, L. macaco, L. albifrons, L. coronatus, L. varius, Nycticebus
tardigradus (3 examples), Perodicticus potto, Galago crassi-
caudata.
Although it is correct to say that there is only a right azygos
in these Lemurs as a complete vein entering the vena cava anterior
on its own side of the body, there are in a few forms traces of the
left azygos—of a hemiazygos. Thus in Galago crassicaudate there
is a longitudinal trunk springing from the right azygos opposite
to the 7th intercostal of the right side of the body. This vessel
runs forward, receiving intercostals from the left side and finally
enters the left subclavian. This vein is obviously the left superior
intercostal of man, but is more extensive in Galago than in Homo.
Furthermore, a posterior fragment of the same vein and not con-
tinuous with it is left in the shape of a longitudinal connection
between the 8th and 9th intercostals of the same side of the body.
In Lemur albifrons I found a hemiazygos arising from the right
azygos shortly after the origin of the first of its intercostal
branches, which crossed to the left side and received the inter-
costal veins of that side of the body.
Of the higher Primates I have examined a considerable number
of species, in all of which there is but one azygos, the right, with,
198 MR. F, E. BEDDARD ON THE [Feb. 19,
in cases, a left hemiazygos. I do not for the present give any
details concerning the Monkeys and Anthropoid Apes.
(5) EpENTATA.
Of this group of mammals I have only been able to examine a
few of the South-American forms.
In the Great Anteater, I/yrmecophaga jubata, there is only a
right azygos, with no traces that I could discover of the left-hand
vessel. As this statement refers to three examples, including
both sexes, it 1s probably a statement of the normal state of affairs
in this animal. In one example, at any rate (I have not notes on
the others), the first affluent occupied the 4th intercostal space.
Of the small Anteater, Zamandua tetradactyla, | have examined
but a single specimen, in which the conditions of the azygos were
quite the same.
In two Armadillos, Dasypus villosus and D. minutus, the right
azygos was also alone present. Hyrtl, in his account of the
anatomy of Chlamydophorus, only found, or at least only men-
tions, the right azygos. These facts afford an additional argu-
ment, though doubtless a small one, for the banding together of
the American Hdentates.
(6) CARNIVORA.
The azygos vein in this order of mammals presents a very
uniform arrangement. I have dissected a considerable number
of species belonging to many genera, and in the great majority of
them there is but one azygos present, which is that belonging to
the right side of the body. In these I have not been able to find
any trace of the vessel of the left side. The following species
present this condition of the azygos veins, viz. :—Galictis barbara,
Crossarchus obscurus, Cynictis levaillanti, Viverra civetta (2 ex-
amples), Vandinia binotata, Cercoleptes caudivolvulus (3 examples),
Nasua rufa (2 examples), Felis pardus, Procyon cancrivorus
(3 examples), P. lotor, Lutra vulgaris (2 examples), Helictis per-
sonata, Cryptoprocta ferox, Herpestes griseus, H. pulverulentus,
Arctogalidia trivirgata, Proteles cristatus, Gulo luscus.
In two specimens of Procyon cancrivorus the azygos of the
right side received a hemiazygos from the left side about halfway
down, and there were no vene intercostales supreme. In Procyon
lotor I found no hemiazygos, but there were two vene intercostales
superior on the right side, supplying the first two ribs and opening.
separately into the precava. On the left side a single vein opened
into the subclavian.
The azygos vein in the Hyena (Hyena crocuta) (text-fig. 68) is
the most remarkable modification of this vein which I observed
in the Carnivora. As has been noted, the azygos in the Carnivora
is constantly a long S-shaped vessel lying on the right side and
extending down to the diaphragm, giving off regular branches. In
1907.] AZYGOS VEINS IN MAMMALS. 199
this Hyena, however, the azygos is singularly short, supplying
only three intercostal spaces. It opens into the Ductus Cuvieri by
a single stout branch ; this is formed by the union of three branches
which are of very unequal calibre. The most anterior of these is
a moderately stout vein running slightly forward in direction.
The middle vein of the three is very markedly the stouter. It
plunges at once, after the shortest possible course, into the thick-
ness of the parietes. The third branch is more slender than both
the first and the second. It also has a very short course obliquely
backwards, and is very soon lost in the parietes. it gives off one
branch to the left side and a slender short main azygos. I could
discover no vein upon the left side of the body. It is to be
remarked that we have here an abortive azygos, the first branches,
or perhaps branch, only being well represented. These collectively
correspond, as I think, to the first branch of the azygos in some
other forms, which is very frequently subdivided into two or even
three branches.
Text-fig. 68.
Azygos of Hyena crocuta. Lettering as in text-fig. 62.
It is usual for the first branch of the azygos to arise between
the fifth and sixth ribs in this order of mammals. I have found
this to be so in Procyon lotor, Lutra vulgaris (one example; in
another it arose between ribs 4 and 5), Swricata tetradactyla,
Genetta felina.
In Crossarchus obscurus (text-fig. 69) the azygos entered the
precaval about opposite to the fifth rib. Its most anterior affluent
arose between the third and fourth ribs. Between the first and
second ribs an intercostal arose on the same side (the right) and
200 MR. F. E, BEDDARD ON THE [ Feb. 19,
Text-fig. 69.
Pc
Scl-= - Sel.
Qy | 7
St mb sa pails
LZ
3
}
al (
Al
“a |
AZ rE
Azygos of Crossarchus obscurus. Lettering as in text-fig. 62.
entered the subclavian vein. The corresponding vein of the left
side of the body was formed of two aftluents arising from the first
two intercostal spaces, which opened into the precaval and noé into
the subclavian.
(7) MarsupPIALs.
T am able to give rather a fuller account of the genera of this
group than of the other groups of mammals treated of in the
present communication. And furthermore, of several genera
which I have examined I have had the opportunity of seeing
several species, and occasionally more than one example of the
same species. The azygos veins of Marsupials have been lately
the object of a careful study by Prof. McClure * in connection
with a detailed survey of the blood system of the Opossum,
Didelphys marsupialis. He has in his memoir collected together
what has been written upon the subject of this vein, and I
shall review his conclusions in the light of the additional facts
which Iam here able to contribute to this branch of anatomy.
* American Journ. Anat. 11. 1903, p. 371.
«
1907. ] AZYGOS VEINS IN MAMMALS. 201
T shall commence with the Kangaroos, of which I have examined
a good many different species.
In Macropus wualabatus the condition of the azygos veins is the
most primitive of that to be seen in Marsupials. There are two
veins, one on each side, each, of course, opening into its corre-
sponding vena cava anterior. These veins appear to be exactly
equal in length.
In Macropus agilis I have found one of the two opposite
extremes. In the single example of this species which I have
dissected there was but one azygos, lying upon the right side of the
vertebral column. In the preceding species of Macropus the two
azygos veins end, or at any rate dwindle to almost nothing, in the
neighbourhood of the last rib behind the diaphragm. Their last
affluent is derived from the body-wall close to the last rib. In
the specimen of JMacropus agilis, on the other hand, the single
right azygos is continued as a wide vein for a long distance back-
wards. <A little in front of the kidney it ceases to be superficial
and is imbedded in the dorsal musculature. It then emerges and
gives off a branch joining the vena cava posterior in the region
of the kidney. This vessel is not the renal vein itself, for the
latter can be followed from the kidney to the vena cava as a
distinct and much thinner vessel. Behind this again the azygos
dies away towards the pelvis, retaining till its extremity a con-
siderable size.
In Macropus melanops I found in one individual the same state
of affairs as in the last species in so far as only a single azygos
was present, and that on the right side. This vein, however, was
slender and drew blood from only six intercostal spaces, after
which it dwindled or entirely disappeared. In a second specimen
of the same species there was also a limited azygos on the right
side ; but, in addition to this, traces of one belonging to the left
side of the body, which, however, was quite small and only drew
blood from one intercostal space.
An example of Wacropus bruni showed conditions intermediate
between the two extremes already dealt with. In this species
there was an azygos on the right side extending as far back as to
the diaphragm. On the left side there was a less completely
developed vein only drawing blood from four intercostal spaces.
A female Macropus dorsalis was not very different. In this
Kangaroo the azygos was only properly developed upon the right
side of the body. It was prolonged, however, for a long way back
as in Macropus agilis. In the same way a branch was given off
to the vena caya posterior underlying the renal vein. The AZY Os
trunk ended some way behind this.
On the left side of the body there were traces of a left azygos.
This consisted of the azygos proper, being a short vessel gathering
blood from one intercostal space only, and of another trunk in
front of this flowing separately into the vena cava anterior of that
side of the body, which corresponds to the superior intercostal of
other mammals.
202 MR. F. BE. BEDDARD ON THE [Feb. 19,
A male Macropus giganteus showed an almost exactly similar
arrangement of the several azygos veins. On the right side the
vein was strongly developed and passed back to the pelvic region,
giving off a branch to the vena cava posterior in the region of the
kidney. On the left side one vein arose from the vena cava
anterior sinistra which apparently corresponds to the two which
arose separately in Macropus dorsalis. This vein was formed of
two afiluents only.
A female Macropus rufus possessed a well-developed azygos
upon the right side, which however did not extend backwards to
anywhere near the pelvic region. I traced it to the eleventh rib,
between which and the tenth lay its terminal branch. The vein
supplied all the intercostal spaces between this and the fifth
anteriorly. On the left side of the body wasa rudimentary azygos
of two branches lying between the fourth, fifth, and sixth ribs.
A second example, a male, showed an identical arrangement of
both azygos veins.
Macropus alligatoris is precisely like Macropus rufus. In a
male specimen of this Kangaroo the well developed azygos lay
upon the right side of the body, while on the left side was a rudi-
mentary vein formed of two afiluents only.
Exactly the same description is to be given of J/acropus
antilopinus.
Of Macropus derbianus I have had the opportunity of seeing
three individuals. Two of these were of particular interest since
they were mother and daughter. The female fcetus showed two
azygos veins very nearly equal in extent, but the right was rather
longer than the left ; this diserepancy—it will be observed—being
the usual one. On the other hand the parent showed reverse
conditions; the left azygos was longer than the right, and in show-
ing this character was unique among the examples of this genus
which I have examined up to the present. But ina third specimen
of the same species, adult and a male, the azygos veins were as
nearly as possible equal, the left being again a trifle the longer.
In Halmaturus bennettii 1 found the azygos on the left side
to be distinctly longer than that of the right side. JI counted in
fact seven affluents on the left side. On the right side there
were only five branches, of which the first followed thie fifth rib. In
a second specimen (text-fig. 70), a male just out cf the pouch, the
small left azygos had only three afiluents from ribs 4 to 6. The
right azygos began with an affluent from rib 2.
“Of the genus Dendr olagus | have examined only one species, viz.
D. bennettii. In this Tree-Kangaroo I found the azygos to be
present only on the left side.
Apyprymnus rufescens is a species which shows differences in
the proportions of the two azygos veins. In one example, a female
the left azygos was shorter, composed of four affluents; the right
reached to the diaphragm. Ina male the right was very much the
shorter, composed as it was of only two affluents. On the left
side there were four or five branches before the vessel disappeared
1907. ] AZYGOS VEINS IN MAMMALS. 203
into the thickness of the musculature. Later on this azygos
dilated to form a thicker vessel which lay beneath the aorta and
reached nearly to the iliacs. This is obviously quite comparable
to what I have described above in Macropus giganteus and other
species; but in them it was the right azygos which was thus
prolonged and enlarged. J could not be certain whether in Zpy-
prymnus rufescens there was or was not a connection with the
renal vein or the postcaval. In this specimen there was also on
the left side a superior intercostal vein flowing separately into the
vena cava superior.
Text-fig. 70.
'Az.F.
Azygos veins of Halmaturus bennettii. Lettering as in text-fig. 62.
In a third example of Wpyprymnus rufescens, which was a
female, the conditions observed differed from those of either of
the other two. The two azygos veins were absolutely symmetrical
so far as I could see, and moreover this symmetry extended to
quite small details. On both sides of the body the last two of the
four intercostal affluents of each azygos joined before pouring
their contents into the azygos. It is curious to find here in the
same species all the chief varieties shown by the azygos.
While Zpyprymnus rufescens is an example of a Marsupial
in which the condition of the azygos veins varies from individual
to individual, the Common Phalanger (7richosurus vulpecula)
offers precisely the reverse characteristic. Of this species I have
dissected six examples belonging to both sexes. In all of them
the azygos vein of the left side is fully developed and reaches back
as far as the diaphragm. In all of them the right azygos is present
204 MR. F. HE. BEDDARD ON THE [Feb. 19,
but is invariably small, and consists at the most of a short trunk
made up of affluents from three intercostal spaces only. There is
some difference in the exact number of these aftluents; there are
either two or three or apparently only one. In one example I
recorded the further extension backwards of the left vena azygos
till it reached the vena cava behind the kidney. This dis-
position of the azygos agrees with what has been already recorded
by previous observers concerning this species. These previous
observations refer to several examples. The state of affairs may
therefore with confidence be regarded as characteristic of the
species.
Allied to the Vulpine Phalanger is the Wombat (Phascolomys
mitchell) (text-fig. 71), of which I have dissected two examples from
Azygos veins of Phascolomys mitchelli.
os. (Hsophageal vein. Other letters as in text-fig. 62.
the present point of view. In the first example, a female, two
azygos veins were present and about equalled each other in length.
The conditions were in fact like those of Macropus ualabatus.
In a second Wombat, also a female, the conditions of the
azygos velus were apparently identical. Both veins were present
and well developed; but the right-hand one was the longer
of the two. In front of the two azygos veins no intercostal
1907.] AZYGOS VEINS IN MAMMALS. 205
branches entered either of the two precavals. The right azygos
entered the precaval at about on a level with the fourth rib. The
first branch arose between the third and fourth ribs. The position
of the opposite azygos was the same. The right azygos continued
down to the tenth rib unaltered; at this point it divided into two
branches, of which the outer supplied the intercostal spaces ; the
inner branch, which was more slender, received at least one cross
branch from the outer and could be traced some little way back as
a very slender vessel; I did not ascertain its posterior connections
ifany. Just after its bifurcation it was crossed by the first inter-
costal artery to cross it, those in front lying below the azygos.
This division of the azygos posteriorly reminds us of the figure of
the Pig’s azygos given by Messrs. Parker and Tozier*. “But it
seems more likely ‘that the inner branch is not the hemiazygos,
but possibly the subcardinal of the right side, the outer branch
being in that case the true persistent postcardinal. These matters,
however, are more fully dealt with below. The left azygos of this
example of Phascolomys mitchelli was shorter than the right; it
ended absolutely between the tenth and eleventh ribs.
In the Brush-tailed Rock-Wallaby (Petrogale penicillata) the
disposition of the azygos veins was like that which is on the
whole characteristic of the genus J/acropus. I have dissected
three examples, one male and two females. They all agreed
except in minutiz. In all of them the right-hand azygos was the
one to be well developed. But I never found that this vessel had
a prominent backward prolongation to the pelvic region as is
occasionally to be seen in Macropus. On the left side there is
only a trace of the azygos, and this is either derived from only
one or from two intercostal twigs. In P. wanthopus, according to
Parsons‘, the right azygos is also the larger, or rather the only
one present.
I do not know whether it is necessary to separate Bettongia
peniciliata from B. ogilbyi. The condition of the azygos veins
offers no help in deciding this question, since the two examples of
B. penicillata differ considerably from each other, and I have only
a single example whych came into my hands labelled Bettongia
ogilbyi. In one specimen of Lettongia penicillata, a female, the
right azygos alone was well developed, extending far back towards
or to the diaphragm. On the left side the azygos was very short,
consisting of two intercostal affuents only. In the second example
of this species, which was also a female, the azygos on both sides of
the body was about equally developed. In the single ettongia
ogilbyi the two azygos veins were well developed; but that at the left
side was distinctly the longer of the two. It is to be noted that if
we are to unite these species, as Mr. Thomas has done in his
‘Catalogue of Marsupials in the British Museum,’ the differences
exhibited in respect of the azygos vein are almost exactly the same
as those shown by “pyprymnus rufescens.
* Bull. Mus. Comp. Zool. xxxi. p. 138, fig. 4
+ See P. Z. S. 1896, p. 706.
206 MR. F. E. BEDDARD ON THE [ Feb. 19,
In a single example of Dromicia nana the azygos was well
developed on the left side of the body and there was a short
azygos on the right side.
In a species of Pseudochirus the azygos was present on the left
side only ; in addition to the azygos an anterior intercostal flowed
separately into the vena cava anterior, deriving its blood from a
single intercostal space only. In a second example (P. peregrinus)
I found the same. The azygos proper commences with the second
rib and joins the vena cava in the neighbourhood of the left kidney.
A single male example of Petawrus breviceps, which was injected
to illustrate the anatomical relations of these veins, possessed an
azygos on the left side only.
In the Thylacine the left azygos is the predominant one, and
there is also a left superior intercostal vein entering the precaval
independently of the azygos. The right azygos is, however, by
no means rudimentary for it supplies four intercostal spaces.
Phascologale penicillata has likewise a well-developed azygos on
the left side extending back to the diaphragm. I saw none on
the right side of the body.
In a female Dasywrus viverrinus the azygos veins were almost
exactly like those of the Thylacine. The left-hand vein was well
developed, extending back almost as far as the diaphragm ; on the
right the azygos was formed of five affluents. Inamale D. mauget
the left azygos was well developed and there was no trace of a
right-hand vein.
A female Perameles obesula agreed with Phascologale rather
than with the last species; for the azygos was present only on
the left side of the body.
Of Onychogale frenata I have examined a single female example.
The azygos was predominantly developed on the left side of the
body. It flowed into the jugular at a point about opposite to
the fifth rib. The first branch of this left azygos formed the
intercostal vein lying between ribs 4 and 5. Thence regular
branches were given off as usual. The main stem of the vein
became much thinner in the region of the diaphragm ; but there-
after increased notably in volume and could be traced back down
into the pelvic region as a massive vein of not much smaller
calibre than the vena cava, alongside and to the left side of which
it ran to a point which I did not determine. There is an obvious
similarity in this case to that of Wpyprymnus rufescens described
above. The azygos was connected to the vena cava by a stout
branch in the neighbourhood of the kidney. An anterior vein, the
superior intercostal, arose separately from the left jugular and
supplied ribs 3 and 4, There flowed into the right jugular
at a point corresponding to that of the left azygos a much
smaller right azygos. This slender vein only drew blood from ribs
4 to 7.
The results obtained from the dissections enumerated in the
foregoing pages enable me to revise some of the conclusions
arrived at by Dr. McClure in his memoir referred to above.
1907.] AZYGOS VEINS IN MAMMALS. 207
Dr. McClure is of opinion that “a single azygos vein is the rule
in Marsupials, and that when two are present the case may be
regarded as a variation.” The facts gathered by myself seem to
me to show that the double azygos of the Monotremata is largely
preserved in the Marsupials, so much so that the process of dis-
appearance of one or the otheris but rarely completed. There are
but few forms in which there is absolutely no vestige of either
right or left azygos as the case may be. This is precisely what
would be expected in view of the admittedly archaic position of
the Marsupials, accepting, of course, the view now generally held
that they are not intermediate between the Prototheria and the
Placentals, but are an offshoot of an early Eutherian. For in
the Rodents, an admittedly primitive type, we have considerable
traces of the double azygos. This latter point, however, will be
discussed in relation with the azygos in Mammalia generally on
a subsequent page. On the other hand, my own observations
confirm Dr. McClure when he remarks that a right azygos is
characteristic of the genus J/acropus, and that a left azygos is
characteristic of the Phalangeride.
J would rather extend Dr. McClure’s remarks about the condition
of the azygos vein in the Carnivorous Marsupials, and point out
that in this whole group the existence of a prevalent left azygos
is the rule so far as we know at present. The table given by
Dr. McClure of the condition of the azygos in such Marsupials as
were known when he wrote, brings out very clearly the variation
in respect of the veins not merely from genus to genus but from
species to species. This generalisation I confirm with great con-
fidence, and also would emphasise the very frequent variation of
these veins from individual to individual. The condition of the
azygos in fact is by no means so fixed in this group as it is in the
Carnivora, for example; and modification is evidently progressing
along two lines, in ene of which the right azygos and in the other
the left azygcs is being retained, while the opposite vein is in
process of disappearance.
Noteworthy, too, is the occasional large size of the postdia-
phragmatic continuation of the azygos in the Kangaroos, Macropus
giganteus, M. agilis, &e., which is, so far as my own observations
go, invariably of the right azygos.
The above results may be conveniently tabulated as follows :—
Azygos on both sides equal or nearly equal.
Macropus wualabatus M. bruni, MW. derbianus, M. bennettit,
Phascolomys mitchelli, Bettongia ogilbyt, Eipypr ymnus rufescens,
Thylacinus cynocephalus, Dasyurus viverrinus.
6. Azygos large on right side, with a rudiment on left of not
more than three intercostal affluents.
Macropus dorsalis, M. giganteus, M. rufus, M. alligatoris,
M. antilopinus, M. melanops, Petrogale penicillata, Bettongia peni-
cillata.
208 MR. F. E. BEDDARD ON THE [ Feb. 19,
c. Azygos only present on right. No rudiment on left side.
Macropus melanotis.
d. Azygos large on left side, with a rudiment only on right
side.
Trichosurus vulpecula, Dromicia nana, Epyprymnus rufescens,
Onychogale frenata.
e. Azygos on left side only. No rudiment on right side.
Dendrolagus bennettii, Didelphys nudicaudata, Pseudochirus,
Perameles obesula, Petaurus breviceps, Phascologale penicillata,
Dasyurus mauger.
(8) RopeEnts.
Of the azygos in this order of Mammalia Owen wrote that
“opposite proportions [to those shown by the azygos veins of the
Hedgehog] prevail in Leporidee and some other Rodents, as in
Squirrels, the left azygos being small or wanting.” ‘This state-
ment is largely but not entirely true, as will be seen in the course
of the following description of several genera and species of
Rodents. In view of the fact that so many Rodents possess, as
is well known, both precaval veins, it might be expected that
here, as among the Marsupials, where a like condition of the pre-
caval exists, there would be traces of both right and left azygos.
And this is precisely what is found among the Rodentia.
Tt is for this reason that I treat of the Rodents next to the
Marsupials.
The most remarkable condition of the azygos vein so far as
concerns the Rodentia is seen in the Beaver. But as I have only
had the opportunity of dissecting a single example of this Rodent,
the structure may be abnormal in the individual. There is only
a single azygos; but the vein, instead of lying upon the right
side, as is otherwise the case when the vein in question is only
developed upon one side of the body, lies upon the left side.
This state of affairs is, however, only an exaggeration of that
found in Jaculus orientalis. In this Rodent, as I have ascertained
and shall point out, of the two azygos veins present that of the
left side is the larger instead of, as in the majority of cases where
there are two azygos and a discrepancy in size between them, the
right being the larger.
This condition of the azygos, however, exists among the Mar-
supials, as has already been pointed out*, and also among the
Ungulates +, but apparently nowhere except as an anomaly.
Of Capromys pilorides I have had the opportunity of studying
two examples which, although they present certain differences,
agree in certain main features in which they also present unmis-
takable points of likeness to their allies Hystrix and Dolichotis.
The azygos is either only or almost entirely to be found on the
right side~. The vein on that side in one specimen follows after
* Supra, p. 202. + Supra, p. 190.
t Dobson (P. Z. 8. 1884, p. 248) only mentions a left azygos.
1907. | AZYGOS VEINS IN MAMMALS. 209
an anterior branch from the right jugular. This latter branch
divides soon after issuing from the jugular (to reverse for the
convenience of description the direction of the blood-flow) into
two, one anterior and the other posterior. Each of these supplies
several intercostal spaces, the anterior two and the posterior
three. The azygos proper does not branch until after it has
passed the region supplied by the branch referred to. Its first
twig supplies two intercostal spaces. The rest are single branches.
On the left side of the body is a single vessel supplying only one
space. In the other specimen I observed no branch on the left
side of the body. On the right side the first branch of the azygos
runs forward and supplies on each side of the body three inter-
costal spaces. It obviously corresponds to the anterior branch of
the jugular of the last individual.
In Celogenys paca 1 have observed rather different arrange-
ments of these veins in two specimens. This animal, unlike
Dolichotis, has two jugulars. In one individual (both were males)
the azygos was developed on both sides of the body though con-
siderably longer on the right. On this side of the body a branch
is given off early in the course of the azygos which supplies
several intercostal spaces; the main trunk of the vein runs of
course parallel with this, and gives off no branches until the last
of those supplied by the large branch referred to. In this speci-
men the inferior intercostal veins do not open independently into
the jugular, but into the azygos before it communicates with the
jugular. In the second individual both right and left superior
intercostals were quite independent of the azygos of thei side,
entering each jugular separately. The azygos vein of the left
side of the body although present was very slightly developed,
apparently collecting blood from only one intercostal space.
In Dolichotis patachonica the conditions of the azygos were as
follows:—The trunk of the right side is the chief one to be
developed. It is, however, as in some other forms (e. g. Wacropus
dorsalis *), derived from two branches which arise separately from
the jugular. The first is a small branch which runs in a forward
direction. The second trunk is the main azygos. ‘This at once
gives off a branch which divides into three twigs, the rest of the
branches are single in their origin from the azygos. On the left
side of the body a vein arises three or four ribs further forward.
Before reaching the parietes it gives off a slender twig which
runs backwards along the carotid and the aorta. I have not
traced this vessel out to its end; it is evidently to be compared
with a similar vessel in Lemur macaco. When it has reached
the parietes this azygos divides into two vessels which run in the
same straight line and are continuous fore and aft along the body-
wall. Two intercostal veins arise from the anterior section and
three from the posterior section. I did not detect any connection
of this left-hand vessel with the subclavian vein anteriorly.
* But in this case it is the left azygos which shows the peculiarity mentioned.
Proc. Zoot. Soc.—1907, No. XIV. 14
210 MR. F, E. BEDDARD ON THE [ Feb. 19,
In another example of Dolichotis patachonica, the conditions
observed were a little different. The azygos proper was only to
be found on the right side of the body, in correspondence with the
fact that this rodent has but one jugular vem. In front of the
azygos are two separately arising superior intercostal veins. On
the left side the vein described in the first specimen of Dolichotis
also existed, but was of much more limited extent than in that
individual. It corresponded in place of origin with the upper of
the two separate superior intercostals of the right side. The
vessel divided into two branches only, of which one was behind
the point of emergence of the vein from the jugular. It is clear
that this trunk is the same as that which in the other specimen
takes up the place of the azygos of the left side of the body, though
it is much less extensive. There is thus no essential difference
in respect of the veins of the intercostal system between the two
individuals. Only a difference of degree in the development of
the same, and that a slight one.
The Porcupine (Hystrix cristata), belonging as it does to the
same subdivision of the Rodents as the Patagonian Cavy, naturally
shows some resemblances to that rodent in the disposition of the
azygos veins. The right-hand trunk is the chief one to be
developed *. In front of it, however, arise from the right jugular
separately two intercostal veins. The main trunk gives off first
a branch which supplies one rib. This is followed by a more
important branch which supplies three intercostal spaces as does.
the corresponding vessel in Dolichotis. After this the branches
are single. On the left side of the body the vena cava anterior
sinistra (the left jugular) gives off two vessels following each other
which arise at about the same level as the two branches of the
right jugular which he in front of the azygos proper. The second
of these is the most important, and runs for some distance down
the body, giving off four branches to the intercostal spaces. All
these lie posteriorly : there is no forward continuation of the vein
such as occurs in Dolichotis; but the forward extension of the
vein in Dolichotis is evidently replaced in Hystrix by the separate
twig arising from the left jugular.
In a second example of Hystria cristata, a male, the arrange-
ment of the azygos veins (text-fig. 72) was apparently identical.
T use the word “apparently” because in the first example I did
not count the ribs between which the various branches of the
azygos ran. In the second specimen the azygos flowed into the
right jugular at a point on a level with the interval between
the third and fourth ribs. The first branch, exactly in the same
way as in the example I have just described, supplied three
intercostal spaces, beginning with the third rib. A superior
intercostal vein arose from the jugular in front of this, and
received from the first and second ribs a branch each. On the
left side the azygos proper was very slender, but supplied ribs
* Parsons (P.Z. S. 1894, p. 685) states that there is “only one azygos vein” in
Atherura africana.
1907. ] AZYGOS VEINS IN MAMMALS. 211
four to seven, and there was in the same way a superior inter-
costal. The constancy of the first branch of the azygos of the
right side which supplies several branches is remarkable. It
seems to me to be possible that this branch is the real post-
cardinal, persistent here to a greater extent than in most other
mammals which I have dissected. In any case its relations are
perfectly consistent with such a view of its nature. I may
mention that in this example I noted an esophageal twig arising
separately from the left jugular not in common with the azygos.
Scl.-<
ry iL)
Qs
Az. l,
Azygos veins of Hystrix cristata.
ces. (Esophageal vein. Other letters as in text-fic. 62.
In the Canadian Porcupine, Hrethizon dorsatum, the jugular,
contrary to what is to be found in Hystria, is a single vessel.
The system of azygos veins is also single and confined to the
right side of the body. There is no trace that I could discover-
of either azygos or superior intercostals on the left side. The
14*
212 MR. F. E. BEDDARD ON THE | Feb. 19,
azygos presents no features of any special interest ; it is like that
of many other animals. In front are two separately arising
superior intercostals. The difference between these veins in the
present genus and in the true Old World Porcupines is one
more justification of the absence of near affinity between those
superficially similar rodents. I examined two specimens of this
species.
Myopotamus coypu has two jugulars, and there is an azygos
vein opening into each at the same level. That of the right side
of the body is the one which is fully developed and runs a long
way down the body. On the left side a slender twig joins the
jugular of that side which appears to emerge from one intercostal
space only. The arrangement of these vessels in the Chinchilla is
the same.
In Lepus europeus, Arctomys marmotta, Cynomys ludovicianus,
Lagomys roylei, there is but one azygos and that on the right side
of the body.
In Sciurus bicolor there are also two jugular vems as in the
species which have just been described. There is also in the same
way but a single azygos vein arising from the right jugular. The
branches of this vein are collected from the series of intercostal
spaces commencing with that lying between ribs 4 and 5.
Seiurus maximus differs from the other Squirrel mentioned in
the present communication in that there is but a single jugular
vein, which is the right-hand one of other Squirrels. Into this
opens the well-developed right azygos vein, which presents no
differences from that of other members of the genus Sciwrus,
except of the smallest detail. Its affluents commence with that
of intercostal space 5/6. Corresponding to this vein, however,
there is a very small left azygos. This vein collects blood only
from ribs 4 and 5, and it opens into the single jugular just a
trifle in front of the point of opening of the large right azygos.
The right azygos, I should add, in both the species that have just
been referred to extends back considerably further than the
last rib.
Of Sciurus vulgaris I have examined two individuals, one a
male and the other a female. The two agreed in every detail of
structure. This species belongs to that group of the rodents which
possess two jugulars. But in spite of this the azygos vein is
single and is present on the right side only. “I did not observe
any superior intercostal vein arising from the jugular in front of
the point where the true azygos springs. I found the same to be
the case with Sciwus palmarum.
In Hydrocherus capybara there are similarly two jugulars and
only a single azygos, which as in Seiwrus is on the right side of
the body. There was nothing to be seen of an azygos on the left
side, which is perhaps remarkable in view of the relationships of
this rodent. This statement moreover applies to two examples of
the species.
In the Jerboa, Jaculus orientalis, a rodent with two jugulars
1907. | AZYGOS VEINS IN MAMMALS. 213
also, there are two azygos veins well developed. I have dissected
two examples, both females, of this species, and find that in both
the left azygos is rather the larger of the two, markedly so in one
specimen. This is a curious reversal of the prevalent arrangement
in Rodents and is reminiscent of conditions found among the
Marsupials. So far as my experience goes this condition of the
azygos veins is unique among the Rodents. It obviously culmi-
nates in the condition observable in the Beaver, where there is
but a single azygos vein and that of the left side. Precisely the
same series is to be met with among the Marsupials.
(9) The Azygos and other Thoracic Veins in the Young of
Myopotamus coypu, and the Homologies of the Azygos in Mammals.
A recent examination of four young specimens of the South
American Myopotamus coypu has furnished facts of great in-
terest in connection with the real nature of the azygos veins in
the Mammalia generally, besides making a contribution to the
anatomy of these veins in that particular rodent. The four
individuals were born dead, but apparently were of full time.
They came into my hands on the following morning, and were in
admirable condition for study. All four agreed excepting in one
detail, and this apparent divergence may have been due to the
difficulty in tracing the smaller branches of veins. In all of these
examples there were three veins in the thoracic region, as well
as—of course—the post-caval. Two veins (text-fig. 73) were
symmetrical with each other, and lay on either side of the ver-
tebral column near to the point of articulation of the ribs. Of
these veins the right-hand one was constantly the larger. Both
opened into the right and left anterior cava respectively. Branches
arising intercostally were seen in both cases. In front of the
point of entrance of this vein into the precava there were no
representatives of the venz intercostales supreme. That is to say,
no veins to which this term might be applied opened separately
into the precava. But as a matter of fact all the intercostal
spaces from the very first backwards had their intercostal veins,
which in the case of the anterior ones passed backwards to join
the main trunk. They joined the main trunk very nearly at its
point of entrance into the precaval vein, and the slightest shifting
of the posterior set of ees would civide oe affluents of
the precaval into two
vena intercostalis suprema, anal a Seon azygos. But is this
vein to be considered an azygos? This point may be deferred
until after description of the third thoracic vein. This vein was
rather larger than either of the paired veins just considered.
It lay in each case on the right side, or—to speak more accurately
—it poured its contents into the precava on the right and not
into the left-hand one of those veins, paired here as in many
other Rodents. The vein really lay rather medianly in position,
running over the centra of the vertebre. Anter iorly it jomed
214 MR. F, E. BEDDARD ON THE [Feb. 19,
the vein of its side which has just been described and opened
in common with it into the precava.
In one at any rate of the Rodents this vein bent distinctly to
the right before effecting this connection, and in another there
was the same bend and (see text-fig. 73) a slender straight vessel
continued on the direction of the vein and apparently opened into
the paired vein rather nearer to the precava. It was generally
to be seen—possibly invariably present though hard to see owing
to deficiencies in the blood—that this vein was connected by cross
vessels with both of the paired vessels already described as running
Text-fig. 73.
Lateral view of the thoracic region of newly-born Iyopotamus coypu, dissected
to show azygos vein (Az.7.) and persistent postcardinal (post.c.).
alongside of the vertebral column on either side. J am inclined
to think that these cross trunks were segmentally arranged in
accordance with the vertebra. This vessel was longer than either
of the paired trunks, and posteriorly the last cross anastomosis
that I observed ‘on the right side was continuous with a longi-
tudinal vessel obviously continuing on the paired vessel of its
side, though a break occurred between the two. This trunk,
receiving affluents in the lumbar region, ended by opening into
the postcava near to the entry into the same of the right renal
vein. Anteriorly the unpaired vessel lay outside the area which
would be occupied by the intercostal arteries (though I did not
1907. | AZYGOS VEINS IN MAMMALS. aa 3)
actually observe the arteries here); later on it could be distinctly
seen to lie within that area. The intercostal arteries could be
observed to lie outside of the vein. We can now consider what is
the nature of these several veins. There seem to me to be only
three possible views, one of which is so improbable that it may be
shortly considered first ot all and then dismissed. This view is
that the paired vessels are two azygos veins which have been
formed in a way not indicated in the specimens from the post-
cardinals, and that the single more median vein represents the
esophageal branch which is at least very commonly developed.
from the azygos and often leaves it near to its connection with the
precaval. If this is the case the vessel must degenerate very much
in maturity, and its numerous CYross connections with the paired
veins are not easy to understand. It seems in fact unlikely that
such unimportant vessels as the visceral branches of the azygos
should have so portentous a beginning and so impotent a conclu-
sion, The other alternatives are mutually exclusive. Hither the
paired veins are the posteardinals and the unpaired vein is a single
azygos, or the single vein is the one remaining postcardinal and
the paired veins are azygos veins which have been derived from
that and another vanished postcardinal. The first of these views
seems to me to be fairly obviously the correct one; and for the
following reasons. In the first place, paired postcardinals and an
unpaired azygos are more likely than the reverse. This is, of
course, not conclusive, for one ev hypothesi postcardinal might
have disappeared. Secondly, the presumed azygos corresponds
with the adult WMyopotamus where there is only one azygos, and,
as is so general with the azygos, it lies partly outside and partly
inside the intercostal arteries. The median position too of the
presumed azygos is like that of undoubted azygos veins, while the
lateral position of the presumed postcardinals is again In consonance
with that view of their nature. In the next place, Dr. McClure *
has pointed out that the azygos arises from the postcardinal in the
Opossum to the median side of that vein, which is precisely the
position occupied by the presumed azygos in the young Myopotanus.
Furthermore, the break between the thoracic and the abdominal
regions of the paired veins is in accord with the view that they
really are the posteardinais ; for such a break has been described
<n the course of the development of those veins.
It is obvious that these facts throw some light upon the azygos
veins in general. It seems highly probable that we shall have to
distinguish in future between real azygos veins or vein and
persistent posteardinals. It is even possible that the true azygos
vein is always a single vein, and that when there are apparently
two-—one on each side—it is a case either of both postcardinals
persisting or of a single azygos with one persistent postcardinal.
That there should be such large differences even in allied mammals
seems surprising; but it is by no means impossible even on
* Toe. cit. (see p. 183).
216 MR. F. E. BEDDARD ON THE [ Feb. 19,
a priori grounds, if—that is to say—we are permitted to use the
analogy of the development of the postcaval. It is perfectly clear
from developmental facts that in the Marsupials generally that
vein has a different origin posteriorly from that which it has.
in the Rabbit, while McClure appears to have shown that in
different individuals of Didelphys marsupialis the extreme end of
that vein originates differently. In man, according to the facts
collected by Hochstetter, the postcaval has as a variation a
different mode of origin from that which is normal.
In the young pouch-living individual of Macropus derbianus de-
scribed above *, it was pointed out that there were two azygos veins
in both foetus and parent. There was no trace in either that I noted
of any other vein. It seems to me to be possible that in that and
other Marsupials there are not any azygos veins at all, but the
persistent cardinals take their place. That this is not the case,
however, in Didelphys appears to be shown by McClure’s investi-
gationst. Again, the relations of the azygos in Cervus sika
described abovet lead one to the inference that the anterior
branch of the azygos running for some distance to the outside of
the azygos is a remnant of the postcardinal of that side, since the
junction of the two veins near to their entrance into the jugular
recalls exactly the conditions which have been described above in
the young Myopotamus. These remarks, however, cannot be
considered at present as more than suggested possibilities. It
seems to me to be equally likely that in those cases where a well-
developed right azygos or left is accompanied by a less developed
vein on the opposite side of the body which I have called azygos
in the preceding pages, the latter is really a persistent post-
cardinal.
In addition to these there are cases where on one side of the
body at any rate both azygos and postcardinal appear to persist.
Those instances present as a permanent condition the state of
affairs which occurs temporarily in Myopotamus coypu. In one of
two Celogenys paca described above, there is, as I regard it, a
well-developed persistent postcardinal on the right side and traces
of the same on the left, in addition to a well-developed ALY OS
on the right and a good but not so well-developed azygos on
the left. In Capromys pilorides there are also considerable
remains of the right postcardinal as well as of the right azygos.
In Hystrix cristata (see text-fig. 72, p. 211) we see precisely the
same thing on the right side; in this species in both examples
dissected. Dolichotis patachonica has also, if my views are correct,
both a right azygos and a short right postcardinal. Nor is this
retention of a feetal condition peculiar to Rodents, though clearly,
so far as my experience goes, most abundant in them; for it also
occurs on the right side of the body in Cervus sika.
The relationship of the posterior end of the azygos to the
postcardinal of the right side requires further emphasis. As has
* Supra, p. 202. + Loe. cit. p. 183. t Supra, p. 192.
1907.] AZYGOS VEINS IN MAMMALS. 217
been mentioned, the azygos bends over the vertebral column and
joins a vein running nearer to the ventral side of the body, which
is hidden posteriorly by the kidney and opens into the post-
caval near to the entry thereinto of the renal vein. Previously
to this the vein has collected tributaries from the lumbar body-
wall. This is precisely what occurs in other and adult mammals.
It is frequently the case that the azygos, after emitting the vem
to the last intercostal space, curves in to the same way over the
vertebral column and can be traced back to the kidney, in the
neighbourhood of which it debouches into the postcaval vein.
It seems to be clear, from the relations of the different sections of
the veins concerned, that the bend to the right (of the right
azygos) is the equivalent of one of the cross branches, by many
of which the azygos is put into communication with the right
postcardinal. Thus the complete azygos in such mammals is
azygos plus one cross communication with the postcardinal plus
the lumbar section of the postcardinal. The azygos, therefore,
has as complex a formation as has the postcaval itself.
This condition of the azygos in many mammals contrasts with
the conditions to be observed among the Marsupials. Here, as
has been pointed out, it is not infrequent for one azygos, either
right or left, to be a vein of very considerable importance, not
merely in the thoracic region but also in the abdominal region.
In various Diprotodont Marsupials, duly recorded in the pre-
ceding pages, I have found a thick vein in the kidney region,
sometimes even extending further back, in fact quite into the
pelvic region. This vein, which in the abdominal region runs
parallel with the posteaval, is connected with that vem by an
anastomosis. Further forwards it is directly continuous with the
azygos vein of its side, which lies in or very nearly in the same
straight line with it. Not unfrequently there is a thinning
before the two veins meet, but still a continuity. This was seen
for example in a specimen of -Zpyprymnus rufescens. In others
there was no such thinning. These cases are, as it appears to
me, to be best explained on the assumption that here the “azygos ”
of the adult is in reality the persistent postcardinal of its side.
Hence the absence of the bend such as occurs in iJ/yopotamus
coypu. The thinning I put down to the “break” which has
already been referred to as occurring between the thoracic and
abdominal parts of the postcardinal in many mammals
There are two other matters for consideration in view of this
suggestion with reference to the differing nature of the azygos in
the adult. Firstly, the variability of these veins becomes of a
different aspect, since it is not one but two distinct veins or pairs
of veins which vary. The actual variability is therefore reducible
in consequence, and to be considered from different points of
view. The variability in the proportions of the two azygos veins
in the Diprotodont Marsupials is, if my suggestions be correct,
obviously not the same thing as the variability in the two veins
which bear the same name among the hollow-horned Ruminants.
218 MR. F. E, BEDDARD ON THE [ Feb. 19,
Broadly speaking, it appears to me that the true azygos is by no
means so variable a vein as are the persistent postcardinals, a
conclusion which is in perfect accord with general principles. A
partially persistent, though obviously decaying structure, is apt
to vary more than a newer structure, which may be increasing
rather than diminishing in importance. Thus we find that
the true right azygos 1s very constant among the Carnivora.
Secondly, there is the fact of the continuation into the abdo-
minal region even as far as the pelvic region of the azygos vein.
In some cases at any rate, possibly in the majority or even all
those animals which (e. g. Carnivora) possess a true azygos vein,
i. €. secondary vein not the persistent postcardinal, that vein is
continuous posteriorly with a longitudinal vein which receives
lumbar branches and opens indirectly or directly into the post-
caval vein. This part of it I regard, for reasons already put
forward, as a portion of the persistent postcardinal. Now it is
precisely among the Diprotodont Marsupials, whose azygos veins
are, as I think, true postcardinals, that we find a very large vein
continuing into the ebdominal and even pelvic region of the
body *. It seems to me that there is some significance in this
fact, and that it tends to support my contention that among
adult mammals the azygos veins fall into two categories.
(10) The Condition of the Azygos and the Classification
of Mammals.
It is clear from the facts here set forth, that the condition of
the azygos vein or veins has a distinct relation to the mutual
affinities of the several groups of Mammalia, but. in quite a
general way. ‘Thus the Lemurs agree with the other Primates in
having only the single right azygos ; and, moreover, it is in these
two groups alone that an hemiazygos is at all prevalent on the
left side, joining the azygos some way before the latter opens into
the right vena cava superior. All the members of the Order
Carnivora agree with each other in the same character; the.
divergences in these three orders or two orders being of the
slightest when compared with some other groups. I do not
infer from this that there is a special relationship between the
Carnivora and the Primates of course. Both groups have, as I
think, independently arrived at the same state of affairs as
regards the azygos vein. But for each group considered alone
the facts at least agree with the general view of their affinities.
It is important to note that the Lemurs agree with the Apes,
and the Arctoid Carnivora with the Ailuroid. No difference in
mode of life seems to have affected this deep-seated character.
The Otter cannot be distinguished in this particular from the
Civet Cat, or the Raccoon from the Suricate.
* Hochstetter, however, declines to recognise as persistent sections cf post-
cardinal Robinson’s description and figures of a left postcaval extension of azygos
(Studies in Anat. Owens Coll. i. 1891, pl. vi. figs. 1, 2, &c.). But it does not
follow that this objection would apply to Macropus.
1907. ] AZYGOS VEINS IN MAMMALS. 219
Again, the cavicorn Ruminants mostly agree with the Suidz in
the entrance of the left azygos into the heart independently of
the remains (if they exist) of the left vena cava anterior, And in
this Moschus and Hydropotes agree with them. This agreement
is not without interest, particular ly when it 1s considered that the
Cervidee may differ in the relations of the azygos: in Cervus sika
the right azygos being the prevalent or only azygos and entering
in the! usual way the right anterior cava. Most zoologists sepa-
rate the solid-horned from the hollow-horned Ruminants, though
to others the distinction between the two groups of Artiodactyles
appears “fanciful.” In any case the Perissodactyles have their
own plan of azvgos structure, which happens to agree with that
of some Cervide, but distinguishes the Suborder from the Pigs
and hollow-horned Ruminants. In connection with the classifi-
cation of the Ungulates, as supported by the disposition of the
azygos veins, the position and number of these veins in Hyrax
are of particular interest. In this ‘“‘subungulate,” admittedly
primitive, and standing nearer to the base of the Ungulate series
than any other living form, except the Elephant, there is as it
were a hesitation to adopt definitely the form of the azygos veins
to be seen in either Artiodactyle or Perissodactyle.
It may also be fairly said that a low position in the series is
indicated by the persistence of both azygos veins. These persist
in Hehidna, and are very generally prevalent in the Marsupials,
the Rodentia, and, though here our knowledge is more deficient,
in the Insectivora. It is the general opinion that of these groups
at least three are primitive groups, and many would think the
same of the Rodentia. The division of the Marsupials into
Diprotodont and Polyprotodont is justified by the condition of
the azygos veins; for in the Diprotodont division there is a much
greater tendency for the two azygos veins to persist than among
the Polyprotodonts. Furthermore, as emphasising the gap that
separates the Marsupials from most of the higher mammals, it is
noteworthy that in them (7. e. the Marsupials) it is the left azygos
which is apt to be predominant and not the right vessel. In the
Order Rodentia also the conditions of the azygos agree with
current views as to their subdivisions. On the whole the
Hystricomorphine Rodents are those im which the two azygos
veins persist to a greater or less extent. An exception to this
statement would appear to be the genus Dipus. But it must be
remembered that the late Dr. Dobson concluded from certain
facts in muscular anatomy that Dipus was not remote from the
Hystricomorpha *.
(11) The Position of the Azygos with reference to the Vertebre.
In dealing with the development of these veins in Mammals,
Hochstetter has dwelt upon the fact that the heart moves back-
wards and with it of course the blood-vessels, including the
* Journ. Anat. Phys. xvii. 1883, p. 177.
220 MR. F. E. BEDDARD ON THE [Feb. 19,
Ductus Cuvieri, attached thereto. The opening of the posterior
cardinal veins into the Ductus Cuvieri thus undergoes an altera-
tion of position with reference to the thoracic vertebre. Thus
Hochstetter found that in the embryo of a rabbit fifteen days
old “ Der Stamm der V. azygos miindet schon in der Hohe des
Zweiten Intercostalraumes in die Hohlvene ihre Seite,’* while
in the adult its position is further back. It is interesting to note
in relation to such developmental facts the varying position of the
- entrance of the azygos into the precava in different families and
genera of adult Mammals. It would seem plain that if the
azygos vein or veins enter the precaval further forward in one form
than in another, the conditions are in that type more primitive.
On the whole it would appear that as a general rule the point
of entrance of the azygos vein is opposite or about opposite to
the fifth rib. It is at any rate so in Macropus, Lutra, Sciurus,
Suricata, Equus, Lemur, &e. It seems to be the general position
among the Carnivora and Primates. I have not met with any
instance of a shifting backwards further than this point, more
than the very smallest. I have noticed occasionally that the
sixth rib instead of the fifth is the first to emit an intercostal to
the azygos. This was the case, for example, with a new-born Ovis
tragelaphus. Now, although Hochstetter and McClure (working
it is true with different types) disagree as to how much exactly
of the embryonic postcardinals persist in the adult as the azygos
vein or veins, there is no doubt that the actual orifice of the
azygos into the precava represents the embouchure of the post-
cardinal into the Ductus Cuvierl. We can, therefore, argue
concerning the shifting position of this point. Dr. McClure has
himself observed + that the left azygos of Didelphys ‘“ opens into
the left precava about opposite the head of the third rib,” while
“the right azygos vein, when present in the adult, opens into
the precava about opposite the head of the second rib.” It cannot
be said, however, that this position of the azygos, which is to be
regarded as a more primitive position, is In any way characteristic
of the Marsupials. It does nevertheless occur in that group, and
1 have found the same in Psewdochirus peregrinus. It is in-
teresting in relation to this matter to find that in the Insectivore
Erinaceus the Ductus Cuvieri on both sides of the body les
opposite to the interval between the second and third ribs. The
Rodents also show to some extent the same anterior position of
the point of entrance of the azygos into the precaval. Thus in
Hystrix cristata the first affluent of the right azygos arises between
the third and fourth ribs. On the other hand, among the Car-
nivora and Primates, as already said, this forward position of the
entrance of the azygos is never (in my experience) to be met
with.
Indeed, the facts stated at length in the preceding pages, which
have just been briefly reviewed, point distinctly to the persistence
* Morph. Jahrb. xx. 1893, p. 573.
+ American Journ. Anat. 1906, p. 185.
1907. | AZYGOS VEINS IN MAMMALS. OAT
of a more primitive position of the Ductus Cuvieri in animals
which possess as a rule two azygos veins, and are thus also
primitive in this peculiarity, and which moreover are usually
assigned on these grounds to a low place in the Mammalian
series. The whole group of facts fit in with each other in perfect
harmony. J believe that these facts are not without importance
in considering the Artiodactyle Ungulates, or rather the hollow-
horned Ruminants and the Pig tribe, which agree in the characters
of the azygos veins.
The right azygos in these animals is always situated far forwards,
beginning even (e. g. in Oryx beatriz) with the second intercostal
space, and although the left and prevalent azygos does not occupy
what is ea hypothesi a primitive position, there are nevertheless
true azygos veins present. The anterior section of the azygos,
which runs forward of the point of opening into the heart, is
perhaps to be regarded as the left precaval otherwise missing.
There is thus some agreement with the suggestions already
made.
(12) The Venw Intercostales Supreme.
These veins, so named by Hochstetter, and often termed inferio1:
intercostals, are superficial veins made up of one or more of the
intercostals anterior to those which unite to form the azygos
vein or veins. ‘They are the superficial superior intercostal veins
of McClure*. As will be gathered from the foregoing pages, these
veins are not invariably pr resent as trunks opening into the vena
cava superior; nor indeed are they invariably present at all. I
presume that in these cases, where no such veins are apparent,
the circulation is effected by ‘the deep series of intercostal veins
lately figured with great elaboration in the Opposum by Dr.
McClure in the paper already referred to. The details of the
occurrence of these veins in various genera of mammals have been
already dealt with in the foregoing pages. From these it will
appear that the veins in question are by no means of universal
occurrence, and that when they do occur they are not always
connected with the precaval vein or veins. In man, for instance,
these veins are described as entering the left and right vene
anonyme respectively. This appears to hold good for the Primates
generally.
But I do not profess in the present communication to have dealt
at any length with the Primates. Among the Carnivora these
veins are evidently not common, and when they do occur some-
times join the precava and sometimes the vena or ven anonyme.
In the Marsupials also the venze intercostales supreme are not
universal, but when they are met with they seem as a rule to
join the precaval or precavals. These veins are much more
general among the Hystricitorm Rodents, where indeed, so far
as my observations go, they are practically universal and join the
precaval. As a rule, too, in this group the veins are paired and
* Amer. Journ. Anat. vol. 11. 1903, p. 371. + Loe. cit. fig. p. 380.
lo
22 ON THE AZYGOS VEINS IN MAMMALS. . [Feb.19,
each precaval has its own separately debouching intercostals.
That this is not at all due to the prevalence among these Rodents
of two precayals is shown by the same constancy of the ven
intercostales supreme among the Artiodactyle Ungulates. The
exceptions that I have met with to the rule that these veins are
present on both sides and open into the single precaval are but
few in number. Concerning other groups the facts at my dis-
posal are so few that I hesitate to attempt any remarks of a
general character.
(13) Conclusions.
Although the facts detailed in the preceding pages, as well as
those made known by the records of others, might be largely
added to, enough appears to me to be known to permit of a few
general observations upon the azygos vein in the Mammalia.
(1) With the exception of the Cetacea the azygos is always a
well-developed vessel of fair size among the Mammalia, and
generally to be found fully developed upon one side of the body
only, and this the right. Its functions appear never to be taken
up by the vena cava inferior, so that 1t is never rudimentary.
(2) There is only a right azygos with no traces of a left—
except rarely as a hemiazygos not reaching the vena cava anterior
—in Carnivora, Lemurs, American Edentates, and Primates.
(3) The existence of two azygos veins characterises several
Marsupials, Rodents, Insectivora, and most Artiodactyles. In the
two first-named groups the existence of the two veins is associated
with the presence of two superior vene cave. It is not, however,
every Rodent with two superior cavee which has also two vene
azygos. But, as it appears, no Rodent with one superior cava
only has more than one vena azygos. The Artiodactyles are
exceptional in that, while possessing only a single anterior vena
cava, there are two azygos veins. But the left azygos generally
opens into the right auricle with or close to the vena cava posterior.
Rarely there is also a connection with the single vena cava
anterior. Where two azygos veins are present it is unusual for
them to be equally developed. Hither the right or the left is
well developed and the other smaller or much reduced.
(4) The existence of a left azygos only characterises a few
forms belonging to the groups Marsupials, Rodents, and Artio-
dactyles. This condition of the azygos is not, like the two which
have been dealt with, characteristic of any large group.
(5) It is to be remarked that, on the whole, the possession of
two azygos veins is associated with a low position.
(6) With certain exceptions (e. g. Hpyprymnus rufescens) the
azygos vein is fairly constant in its characteristics for a given
species. This is sometimes very markedly the case.
(7) As will be gathered from the details given, the azygos is
of some use in the classification of Mammals. Thus Hyrax shows
its Ungulate affinities, and several genera of Hystriciform Rodents
agree together in the character of the azygos veins, &e.
1907. ] ON THE ORIGIN OF FLIGHT. DONS
(8) As a general rule the entrance of the azygos vein or veins
into the precaval or precavals is more anterior in position in
Mammals, occupying a lower position in the series than in more
specialised types, 7. €. opposite to second rib instead of fifth or
sixth. This corresponds with ontogenetic shifting back of heart
and blood-vessels.
(9) The conditions observable in the newly-born young of
Myopotamus coypu seem to show that the posteardinals may
persist as such at least up to the time of birth, and in some adult
Rodents one is also persistent.
(10) The same species shows that the azygos of the adult is
independent—except for a very short tract at its opening into the
precava—of the postcardinal of its side, thus confirming the embryo-
logical results of others who have affirmed that only the very
commencement of the azygos is traceable to the persistent post-
cardinal of its side.
(11) It is probable that the veins called “azygos” in adult
Mammals are not in every case strictly homologous veins. Where
there is but one azygos present (e. gy. Carnivora) it is probable
that that vein is the true azygos, except in the abdominal region
where it is formed by the persistent postcardinal. In cases where
there are two azygos veins both may be (? certain Marsupials)
persistent postcardinals, or one of the two may be a remnant of
the posteardinal, the other being a true azygos.
(12) These and some other facts and conclusions lead to the
inference that the true azygos vein of Mammals (7. ¢. that formed
by an outgrowth of the postcardinal) is a structure which has been
developed in the Eutheria.
5. Ideas on the Origin of Flight.
By Dr. Baron Francis Nopcsa.
[Received February 8, 1907.]
(Text-figures 74-82.)
Although much has been written on the origin of flight,
yet till now no really satisfactory explanation for this kind of
locomotion has been found. ‘This is, so far as I can under-
stand, mainly due to the fact that it has on @ priori grounds
been supposed that all the principal groups of flying vertebrates—
namely, Pterosaurs, Bats, and Birds—originated in a similar
manner, without fully appreciating the fundamental fact that,
from the mechanical standpoint, patagium and feather are two
perfectly different organs.
A patagium is a soft flexible membrane and in consequence
requires, to be effective, numerous firm radial supports originating
from the body that has to be carried, whereas for a series of
semirigid but elastic quills one line of attachment is sufficient.
In consequence of this difference, a patagium-flyer must always
224 BARON NOPCSA ON THE [ Feb. 19,
adapt fore and hind limbs and tail to the support of the patagium,
whereas in a generalised feathered animal only the feather-
supporting elements need become affected by violent specialisation.
The development of the posterior limb in such an animal is but
little, if at all, affected by the development of flight (foot in Eagle,
Parrot, Woodpecker, Nightingale, Goose, Stork, Ostrich, &c.).
As to flight itself we have to distinguish, as partially already
pointed out by Dollo, three distinct stages of evolution : first
parachute or passive flight, then flight by flapping the wings or
Night by force, and lastly soaring or flight by skill.
As Langley and Lucas pointed out in their highly interesting
papers, the soaring birds lack carrying power (in accordance with
which fact the crista sterni is often comparatively feebly
developed), while flight by flapping of the wings, as shown by
the generally soaring Eagle when carrying prey, enables the
animal to support a good deal of weight.
That soaring birds show a sharply pointed wing, while birds
that fly mainly by flapping display a wing with a more or less
rounded outline, is well known.
After these preliminary, but I think essential, observations,
T shall now point out some characters of Pterosaurs, flying
Mammals, Dinosaurs, and Birds that have not yet been brought
together.
PTEROSAURS.
The Dimorphodon, till now the -earliest-described long-tailed
Text-fig. 74.
Hind limb of Dimorphodon
Pterosaur, shows ‘in its hind limb no sign whatever of cursorial
locomotion (text-fig.74). The metatarsals 1-4 are equally developed,
1907. | ORIGIN OF FLIGHT. 225
but the 5th is somewhat thicker and also much shorter. The
elongate phalanges of the 5th toe further prove clearly that no
cursorial adaptation modified the form of these bones. Very
much the same type of foot is visible in the equally long-tailed
Campylognathus; and when we turn to the Rhamphorhynchus of
the Solenhofen Slate (text-fig. 75), we find not only no cursorial
modification of the four-toed slender foot but quite decided degene-
ration. However, according to Zittel, the number of phalanges in
the 5th toe is perhaps somewhat greater than in the drawings
given for Dimorphodon or Campylognathus. Since the spur-like
clawless 5th digit of the foot is very strongly developed in Dimor-
phodon, there is, as Owen observed, good reason to believe that a
Text-fig. 75.
Hind limbs of Rhamphorhynchus.
uropatagium was not only present but even very well developed ;
whereas we know that in Rhamphorhynchus, im accordance with
the less developed 5th toe, no uropatagium extended to this part of
the body. The resemblance of the Rhamphorhynchus sternum
to that of the Bat (Z’aphozous) likewise has to be noticed.
That in the Liassie Dimorphodon the wing-finger is relatively
shorter than in the Tithonian Rhamphorhynchus is a fact so-
obvious as scarcely to demand attention. The short-tailed
Pterosauria of the genus Pterodactylus, with comparatively short
wing-bones, resemble Campylognathus in having four feeble and
Proc. Zoo. Soc,— 1907, No, XV. 15
226 BARON NOPCSA ON THE [Feb. 19,
equally strong metatarsals which all approximately attain the same
length, while, in harmony with the lack of a tail, of the 5th toe
only a rudiment (text-fig. 76) now remains. Among Chiroptera we
find that in the tailless forms, notably Péeropus, quite similarly
the os calear of the hind leg is less developed than in the long-
tailed species.
Hind limb of Péerodactylus.
(The rudiment of the 5th toe is unfortunately not shown in this drawing.)
On account of the anterior prolongation of the ilium in
Pterosaurs, and on account of the great number of vertebre united
in the sacrum, it has been frequently assumed that the Pterosaurs
enjoyed a bipedal locomotion. Both these arguments, however,
fail to convince me, and this principally on account of Vyctosaurus,
which, although certainly not a bipedal genus, has a still greater
number of sacral vertebre, and because in Pteropus there is like-
wise a pseudosacrum present. Another argument that can be
brought forward as annulling the hypothesis just mentioned
consists in the fact that the Pterosaur pelvis, though showing
considerable length, has an ilium of an exceedingly low and
narrow Bat-like outline.
A Pterosaur of whose crawling habits we can be quite sure is,
as just mentioned, the Upper Cretaceous Vyctodactylus, for, as
Williston pointed out, the acetabulum is placed far back, nearly over
the edge of the sacrum, so that it was impossible for the knees in this
animal to meet in the middle, and at times the knees may even
have been turned more or less backward. When the femora
were rotated outward and abducted, the tibie might have been
brought parallel with each other. Exactly similar conditions are
1907. | ORIGIN OF FLIGHT. 227
to be met with among Bats, whose crawling locomotion is familiar
to every student.
The elongation and attenuation of the hind feet in Vyctodactylus
(text-fig. 77) are also characters that demand mention, and a
similar elongation is again to be met with in the tailless Vampires.
An interesting feature is the co-ossification of numerous dorsals
in Vyctodactylus and the nearly allied European genus Ornitho-
cheirus.
Text-fig. 77.
Hind limb of Nyctodactylus.
(Photograph of specimen in the British Museum.)
A Triassic long-tailed Pterosaurian, Zribelesodon (the detailed
description of which I intend publishing on some other occasion),
shows much the same proportion between total length of hind
and fore limbs as does Galeopithecus ; and, although Galeopithecus
proves to be in no way related to Chiroptera, still we must suppose
that the whole order Chiroptera, considering the patagium, passed
through a Péeromys- and a Galeopithecus-like stage in the course of
its evolution.
Since, as already pointed out, a patagium requires many spear-
like supports, and since in arboreal animals fore and hind limbs
are to the same extent used for running and leaping, it is evident
that primarily fore and hind limbs must have become to the same
extent used for the support of the patagium, which necessarily had
to take origin at the centre of gravity between humerus and femur.
As soon as such a potentially flying animal became actively
228 BARON NOPCSA ON THE [ Feb. 19,
volant, and began to fly by force—that is, move its patagium,
—the sternum must evidently have developed a sternal crest,
the patagium must have continued to increase its surface, and
this then would not only produce stretching of the limbs but also
development of secondary supports of the patagium.
Such secondary supports are, as we shall see, developed at
different times and in different ways, being produced by special
development of the olecranon, the carpal bones, and _ossified
tendons. In the long-tailed Pterosaurs such supports are absent
in the fore limb; in the short-tailed Pterosaurs, however, they are
well developed and are represented by a modified carpal which,
according to Williston, shows (Pterodactylus — Nyctodactylus)
progressive evolution. Another modification that each patagium
produces in the animal’s body is to bring all the radial supports
to the same level, and this, making the acetabulum and knee
rotate outward and backward, produces subsequently crawling
locomotion.
When the last stage of development is attained and aerial
locomotion accomplished by skill and not by force (Wyctodactylus,
Rhamphorhynchus), the patagium obviously not only would assume
a pointed outline and become reduced to a smaller surface, but in
some cases also the tail would change to a rudder-like organ
(Rhamphorhynchus) or become entirely lost, while the attenuated
feet would in this case assume the function of steering (Vycto-
saurus). It is of no small importance that of the two highly
specialised groups of Pterosaurians(Rhamphorhynchide and Nycto-
sauride) the tailless ones should have survived the longer.
MAMMALS,
Very much the same changes as are to be found between
Tribelesodon, Dimorphodon, and khamphorhynchus on the one
hand, and Péerodactylus and Nyctodactylus on the other, are also
to be observed when we come to consider the patagium-flying
Mammals. A set of good diagrams of flying mammals has recently
been published by R. 8. Lull. Petawrus and all other animals
with a small patagium represent the stage where, as in all arboreal
animals, a very long tail is present.
Only a plagiopatagium is present in Petawrus, a propatagiam
is added in Péeromys, whilst in Anomalurus even a uropatagium
is present. Asin Pterosaurs, supplementary patagial supports are
frequently developed. In Anomalurus and Vespertilio such a
support arises from the olecranon, in Pteromys it is partially
attached to the pisiforme and partially, though to a less extent,
to the 5th metacarpal; and in embryos of Chiroptera quite a
similar structure is met with: a modification recalling the back-
wardly directed toe of the hind leg in Dimorphodon is produced
by the development of the caleaneum’s calear projection. As in
Pterosaurs, so also in flying Mammals a very low ilium is present,
and this not only in: Chiroptera but also in Galeopithecus, where,
1907. ] ORIGIN OF FLIGHT. 229
even during ontogenetical evolution, a backward rotation of the
ischium, and in consequence a flattening of the pelvic girdle, is
to be met with. Pteromys and Anomalurus, according to Dollo,
have to be termed passive flyers: the first partially active fiyer
seems to be Gialeopithecus, for, according to Wallace, this animal
is not only capable of sailing downward, but at the end of its
downward oblique glide to rise a little upward.
Galeopithecus, however, is a long-tailed, comparatively short-
armed patagial animal, in which never theless the patagium ex-
tends even to the tips of the digits and to the end of the tail ;
while when we turn from this 0) the specialised actively flying
Chiroptera, we are impressed firstly by the elongation of the
wing, and secondly by the frequent partial or total loss of the
tail.
Both in Pterosaurs and Bats the main movement during flight
seems to have been, and still is, dependent on the humeral articu-
lation. The similarity of ‘the patagial structure in Lhampho-
rhynchus and Bats, as remarked by Zittel, is also to be noted.
The hairless condition of the patagium in ‘Chiroptera compared
with Galeopithecus is likewise a more specialised feature ; while
Pteropus vula SG (more specialised than any Bat in regard to the
caudal region, ‘‘chevauchement de spécialisation ”) * shows, by
possessing some hair on the interior surface of the patagial
membrane, an intermediate stage.
In Pterosaurs, as also in Péeropus, the number of sacrals is
augmented, and in the latter they even form, by co-ossifying with
the ischium, a pseudosacrum.
The more or less perfect reduction of tibia and ulna is another
character that is noteworthy in all patagium-bearing Mammals.
in an analogous manner to the Cretaceous Ornithocheirus, also
in some Bats a rigid thorax is attained, though in this case the
ribs and not the vertebrze co-ossify.
Since we may safely assume that Bats descended from Mammals
which possessed a well-developed neural spine, the reduction of
this process, noticeable also in the Flying Lizard (Draco volans),
has also to be considered as a sign of specialisation
The thin and dense skull-bones also unite in specialised Bats,
very much as in Birds and Pterosaurs; and as to the brain,
there exists a great amount of resemblance between the cast of
the brain-cavity in some Kocene Bats, in Hesperornis, and in
Scaphognathus.
Only in one point is there a pronounced difference between the
Pterosaurs and the Bats, and this is in the transformation of the
phalanges of the pes and manus. While in the Pterosaurs a reduc-
tion of the claws takes place in the pes, and they remain present
in the manus, in Chiroptera exactly the opposite happens; but
this divergence is easily understood when we consider that the
Chiroptera had, in consequence of adapting four fingers to flight,
only their hind feet at their disposition, for resting ‘and suspend-
ing on branches, while the Pterosaurs, which developed only one
230 BARON NOPCSA ON THE [ Feb. 19,
wing-finger, could always suspend themselves by the remaining
free digits. This is, perhaps, also the reason why, in both groups,
ulna and radius, tibia and fibula have been reduced in a different
manner. In Birds the same problem has been solved im quite
another manner (musculus ambiens and peculiarities in the
structure of the tendon-sheath of musc. flexor. digit. im the
phalanges). Text-fig. 78 shows the hind limb of a Bat identified
by Dr. K. Andersen as Hipposiderus gigas.
Text-fig. 78.
Hind limb of Hipposiderus.
Since all flying animals must needs have developed from agile
quick-moving animals, since all living patagium-flying animals
(such as Draco volans, and the other living animals mentioned
in this paper) are arboreal, leaping, quadrupedal creatures, and
since, further, a bipedal cursorial animal, on account of mechanical
impossibilities, can never develop a patagium—for such an organ
would in bipedal (7. ¢., erect) locomotion only catch the air and so
prevent running without raising the body,—and since the union
of fore and hind limbs is directly opposed to bipedal cursorial
locomotion, we can safely state that all patagium-flying animals
originated from quadrupedal, leaping, arboreal forms.
Bats and Pterosaurs, though they support the wing in different
ways, still show an analogous direction of evolution—as shown by
the development of a patagium with all that this implies; thus we
may safely state that Bats and Pterosaurs have arisen in similar
manner from quadrupedal arboreal forms.
1907. j ORIGIN OF FLIGHT. 231
DINOSAURS.
In consequence of the quite extraordinary tendency of Dinosaurs
to specialise every now and then along Avian lines, and in
consequence of the fact that the most primitive Dinosaurs are
bipedal in their habits, it is not only probable that all Dinosaurs
originated from bipedal forms (I only need to quote the numerous
bipedal tracks in the Red Triassic Sandstone in Connecticut), but
that they also are very nearly related to the primitive Birds.
Since Dr. Holland thinks that the Dinosaur likeness to Birds
is sometimes greatly exaggerated, I would like to mention some of
the most characteristic primitive and adaptive Avian features of
Dinosaurian reptiles: basis cranii (Hypsilophodon, Compsognathus),
development of beak (Orthopoda, beak perhaps developed
independently in different suborders, caused by latent homoplasy),
lack of neural spines in cervical vertebrae (Sauropoda), dorsal
neural spines bifid (Diplodocus), saddle-shaped articulating surface
of sacrals (Streptospondylus), synsacrum (Orthopoda), 'pyornis-
like caudals (Diplodocus), Avian scapula (Orthopoda, Theropoda),
co-ossification with the coracoid (all Dinosaurs), manus (Orni-
tholestes), ilium covering last ribs (Sauropoda), ilium touching
neural spines (Stegosauride), ilium showing antitrochanteric
ridge and dorsal plane (Theropoda) ; backward rotation of pubis
and subsequent development of processus pseudopectinealis
(Orthopoda), femur shorter than tibia (many Dinosaurs) ;
reduction of fourth trochanter (all Dinosaurs), distal end of
femur (Streptospondylus, &c.), development of processus ascendens
astragali (Theropoda), fusion of caleaneum and astragalus with
tibia “(Compsognathus) ; position of hallux (Theropeda) ; pheu-
maticity or light structure of the whole skeleton (many Dinosaurs).
In a paper on the evolution of Dinosaurs, I pointed out that
the Theropoda specialise by developing an interpubic ossification,
by augmenting the number of their sacrals,; by changing the
character of their vertebre from biconcave to opisthoccelous, by
lengthening their neural spines in the dorsal region, and by
developing a proc. asc. astrag. and reducing the number of their
toes. In more specialised Theropoda the metatarsals become
always more closely applied, and. lastly, these animals specialise
by losing the fourth trochanter. Most of these changes are also
notable among the bipedal Orthopoda, and since this develop-
ment is independent of that in the Theropoda, we must consider
them as homodynamic changes; besides this, in Orthopoda we
can trace a thickening of the bony matter and the development of
a processus pseudopectinealis. A functionally analogous osseous
process is developed in most running birds after the co-ossification
of the pelvic elements.
Since we can be sure that in Dinosaurs all the changes
mentioned are not due to the giving up of volant habits, but are
merely signs of cursorial adaptation, we have a clue to under-
stand some of the changes that occur among the Paleognathous
Birds. Besides this we can fix the fact that the Dinosaurs, like
232 BARON NOPCSA ON THE [Feb. 19,
many cursorial Mammals, were not only set the problem of
developing a flexible dorsal vertebral column, which was attained
by development of convexo-concave intervertebral articulation,
but that to this, in consequence of the position of their head, they
had to add strength in the vertical direction, which could only be
attained by developing the attachment surfaces for the musculus
longissimus dorsi on the neural spines and the producing of
hypapophysis-like knobs on the cervicals. This, moreover, is the
first consideration adduced since 1887 that shows us that the
vertebral column of Zgwanodon, though provided with ossified
tendons, cannot have been altegether rigid. Text-fig. 79 is in-
tended to show the highly modified foot of the Cretaceous Dinosaur
Ornithomimus, and can be compared with the feet of Dipws and
Alactaga (text-fig. 80).
Text-fig. 79. Text-fig. 80.
J
Text-fig. 79.—Hind limb of Ornithomimus.
80.—Hind limbs of Dipus (left) and Alactaga (right
BirDs.
Leaving Dinosaurs and turning to Birds, we observe the follow-
ing salient points :-—
The first and most primitive Bird we know, Archwopterya
(text-fig. 81), shows not only a perfectly bird-hke femur and tibia,
but also tridactylism, and this is, as demonstrated by Dinosaurs
and the Dipus-like rodents, a prominent feature of bipedal
cursorial or saltatorial specialisation, while it never occurs among
arboreal forms.
The pelvis of Archwopteryx, moreover, is essentially that of a Bird,
1907. ] ORIGIN OF FLIGHT. 233
and as a sign of cursorial locomotion there is even an indication
ef a processus pectinealis. The vertebre are free, and neural
spines are present.
Besides this, Archeopteryx differs from all Birds by having a
long laterally feathered tail, that to a certain degree reminds us
of the peculiarly covered and flat-looking tails in the mammals
Acrobates and Ptilocerus low.
Text-fig. 81.
eae (i oopkese eee PS
Hind limb of Archeopteryx.
(Photograph of the British Museum type specimen.)
The ossified tendons which occur in the tail of Archeopteryx
show further that strength of this organ was required just
as much as in the tail of Dimorphodon or Rhamphorhynchus.
A long tail, sometimes even with ossified tendons, is quite a marked
feature of the Dinosaurian bipedal reptiles, and its loss, as shown
in Pterosaurs, is generallyin harmony with the better adaptation to
flying locomotion.
The rounded contour of the Archwopteryx-wing, together with
the feebly developed sternum, show us that Archwopteryx, though
perhaps not an altogether badly flying creature, can on no account
have been a soaring bird, but a bird that was yet in the first
stage of active flight.
That the soaring Frigate-Birds and Albatrosses have a com-
paratively weaker sternum than the Gallinaceous Birds has already
been mentioned ; and I therefore need only point to the formation
of a rigid thorax in flying birds as analogous to the condition in
Pterosaurians and Bats and in opposition to the Ratite, and to
the fact that the cursorial Paleognathz, contrary to the flying
Proc. Zoou. Soc.—1907, No. XVI. 16
234 BARON NOPCSA ON THE [ Feb. 19,
Tinamous, possessed not only free vertebrve, but even elongate,
Dinosaurian-like neural spines in the dorsal region, and this
because also in this case, for running, strength had to be united
with mobility in the dorsal region, whereas for flight, as already
mentioned, strength and rigidity seem to be the qualities required,
go that the neural spines become to a certain extent useless. It
is especially to the curious dorsal and caudal vertebrae of Apyornis
that I should like to draw attention. Probably mobility is one
of the reasons why in the flightless Hesperornis saddle-shaped
vertebre were developed at a period when Jchthyornis still showed
biconcave articulation, although I am quite aware that perhaps
other explanations will have to be sought for, since also in other
ways Hesperornis indicates a more specialised form, and this not
only by its wing-bones being already reduced, but by exhibiting a
certain tendency to lose its teeth, since these are no longer placed
in distinct sockets as in Archewopteryx and Ichthyornis, but in a
furrow.
If we, after these preliminaries, now suppose that Birds, betore
attaining the Archwopterya-state, originated from quadrupedal
arboreal animals and only after having learnt to fly became bi-
pedal, it is difficult to understand why they in general show Dino-
saurian affinities, why they did not use both hind and fore limbs
to the same extent for flight as they would have done for arboreal
locomotion, why the bones of the pectoral region and of the wings
show more primitive traces than the hind parts of the body, and
why they did not, like all other quadrupedal flying animals,
develop a patagium ; whereas, if we consider that in Archwopteryx
the anterior extremities, though bearing the most important
ectodermal pinions, are less modified than the posterior extremities,
which are already perfectly bird-like, and if we then suppose that
Birds originated from bipedal Dinosaur-like Reptiles, it is easy to
understand what induced the Birds to attain an Archwopteryx-
like stage of evolution, for at first a certain amount of bipedal,
and only afterwards a volant, modification would be required.
While we can safely state that a bipedal animal never could or
did develop a patagium without giving up bipedalism, this cannot
be said of feather-bearing forms, for we may quite well suppose
that birds originated from bipedal long-tailed cursorial reptiles
which during running oared along in the air by flapping thew
free anterior extremities. It Dinosaurs had bird-like pulmonary
appendages, as indicated by the pneumaticity of the skeleton, such
movement would only have been of advantage for the respiratory
organs (the pneumatic foramen occurring sometimes in Moa-bones
would therefore be an atavistic feature, and the loss of pneumaticity
would be a parallel to the same change in the Dinosaurian sub-
class). At this point the pulmonary appendages of Chameleons
have also to be taken into consideration. A double running and
flapping action would—somewhat in accordance with Pycraft’s
views on this subject—subsequently easily lead to an enlargement
of the posterior marginal scales of the antibrachium, and at the
same time produce a certain amount of bipedal specialisation.
By gradually increasing in size, the enlarged but perhaps still
horny hypothetical scales of the antibrachial margin would
1907. ] ORIGIN OF FLIGHT. 235
in time enable the yet carnivorous and cursorial ancestor of
Birds to take long strides or leaps, much in the manner of a
domesticated Goose or of a Stork when starting, and ultimately
develop to actual feathers; this epidermic cover would also raise
the temperature of the body, and thus help to increase the
mental and bodily activity of these rapacious forms. The
possibility of such a development of flight is clearly shown by
the somewhat analogous, but still more marvellous and nearly
paradoxical, yet not unfrequent, development of Flying- Fishes.
The marginal scales being originally the principal wing-element
in such a hypothetical form, these parts could attain quite a
considerable size without essentially altering the underlying bones
of the arm, a fusion of the carpal phalanges being only then
necessary, when in flight rigidity of this region became requisite.
Besides this, the continued use of the anterior flapping limbs as
Text-fig. 82.
Hypothetical reconstruction of a running “ Pro-Avis.”
grasping-organs would also account for the feeble specialisation of
the digits in the Ornitholestes-like manus of Archwopteryx, and
for the preservation of the claws in the Ostrich and Opisthocomus,
where, according to Pycraft, the temporary delay in the growth of
the distal pinions has been developed simply not to prevent the
claws from performing their still not unimportant function. An
effort to condense these hypothetical changes into a drawing is
given in text-fig. 82, which might in consequent allusion to
Pyeraft’s analogous reconstruction be called a “ Pro-Avis.”
The facts that even from the Hocene formation in most parts
of the world numerous big Ratites are known, which can only
have originated from badly-flying ground-birds, whereas in more
modern times the Ratites are apparently vanishing from the
earth’s surface, likewise find quite an easy explanation in the
hypothesis that in the Mesozoic times badly-flying ground-birds,
and not tree-birds, were the prevailing forms. The individual
or ontogenetical development of every cursorial Carimate (for
example, every gallinaceous bird) would thus show us the exact
manner in which flight has been acquired. The true phylogenetic
value of the surviving ‘‘ Paleeognathee,” with their body-temperature
236 ON THE ORIGIN OF FLIGHT. [Feb. 19,
decidedly lower than in other birds on the one hand (Sutherland),
and their reduced brachiosternal muscles on the other (Fiirbringer),
can likewise be appreciated only if we consider them as forms
that specialised at a very early stage of Avian evolution.
It is to be remarked that among the terrestrial birds which
according to this hypothesis would seem to have preserved their
original mode of living and manner of breeding, the nest-building
faculty is less developed than in those birds which, to avoid
the dangers of ground-life, migrated up into the trees and had
then to shelter their eggs and young ones from the new chance
of falling to the ground. That ground-life involves for a bird
more dangers than life on a tree, I think, is shown by the fact
that the true ground-birds usually are protectively coloured, while
in the latter, even among Gallinaceous Birds, bright—one might
nearly say artistic—sexual characters are frequently developed.
The supposition, that Birds once possessed a patagium and only
afterwards developed feathers, I consider as devoid of foundation,
for apart from the impossibility of a marginal feather being
effective, when only attached to a flexible membrane, it is loss
and not development of hair and scales (= epidermal coverings)
that takes place in the Chiropterygian and Pterosaurian patagium.
Besides, I do not see any reason why a useful patagium, once
developed, should suddenly have stopped growing.
The long tail in Archwopteryx can in no way be invoked in
favour of a primitive arboreal stage of Birds, for a long tail not
only characterises arboreal but also bipedal cursorial and saltatorial
forms. Thus we cannot find a single character in Archeopteryx that
would absolutely prove arboreal specialisation, while the develop-
ment of the cannon-bone alone is sufficient to show with certainty
that some of the direct ancestors of Archwopteryx had cursorial
habits.
CoNCLUSION,
From a consideration of the whole of the above remarks, we
can, I believe, formulate the following statement :—
While Pterosaurs and Bats originated independently from
quadrupedal arboreal forms in which both anterior and posterior
extremities, in consequence of the development of a patagiwm,
hecame primarily equally used for flight and in consequence equally
unfit for locomotion on the ground, Birds originated from bipedal
Dinosaur-like running forms in which the anterior extremities, on
account of flapping movements, gradually turned to wings without
thereby affecting terrestrial locomotion. This 2s also the reason why
Birds became dominant over all the rest of their aerial rivals.
In conclusion, I take pleasure in thanking once more all the
ventlemen that helped me to compile this paper, notably Dr. K.
Andersen, Mr. G. A. Boulenger, Dr. Forsyth Major, Mr. W. P.
Pyeraft, and Dr. A. 8. Woodward, at the British Museum of
Natural History.
P.Z.S. 1906, pp. 759-1052 were published on April 11th, 1907.
No. 38.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGICAL SOCIETY OF LONDON
January 15th, 1907.
Dr. J. Rost Braprorp, F.R.S., Vice-President, in the Chair.
The Secretary read a report on the additions that had been
made to the Society’s Menagerie during the months of November
and December 1906.
Mr. Ouprretp THomas, F.R.S., exhibited the skin of a new
Monkey from the Ituri Forest, obtained during the recent Ruwen-
zori Expedition, and diagnosed it as follows :—
CERCOPITHECUS DENTI, sp, n.
Allied to C. campbell, but not darkened on the posterior back
and hind limbs, the outer side of these latter being grizzled olive-
yellowish to the ankles. Under surface and inner side of limbs
very sharply defined creamy white. Tail greyish white, darkening
terminally to black.
Hab. Ituri Forest.
Type. Male. Original number 184. Collected by R. E. Dent.
Mr. P. H. Banr, F.Z.8., read a paper “On the ‘ Bleating’ or
‘Drumming’ of the Snipe (Gallinago celestis).” The object of
the paper was to prove that this phenomenon was produced by
the tail-feathers of this species, a point which had been much
disputed. It was found that if the feathers were attached to a
cork in a special manner, the peculiar bleating sound could be
produced, and, furthermore, that only two feathers in this species
were the active agents in producing the sound. Observation
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance.
2
proved that these two feathers were held in a particular manner
in front of the others during the bird’s flight in the breeding-
season. Feathers of both male and female were found to bleat, a
fact which had been borne out by numerous observers in the field.
These feathers were found to have a peculiar structure, differing
materially from the other feathers in the tail. Microscopically
they differed, and the number of hamuli were found to be in excess
of those found in other feathers. The feathers of various exotic
species had been experimented upon, and those of G. delicata,
nobilis, frenata, paraguaye in the New World, G. australis and
aucklandica in the Antipodes, and G@. solitaria and megala in
Asia had been found to produce musical sounds. These feathers
varied in structure, and consequently the sound produced differed
accordingly. The feathers of G. gallinula, G. major, and G. stenura
were not found to be musical.
Mr. J. L. Bonnote, F.Z.S., communicated a paper on a collec-
tion of Mammals from Annam sent home by Dr. Vassal. Twenty-
four species were enumerated, of which the following four were
described as new :—
1. NYCTICEBUS PYGMUS, sp. n.
Similar in general colouring to, but about half the size of,
WV. coucang, and without any dark ‘markings round the eyes or
down the back. The teeth are quite different, the second molar
being the largest, whilst the third molar is triangular in shape
and but little inferior in size to the first.
2. TUPAIA CONCOLOR, sp. Nn.
Similar in general colouring to 7’. belangeri, but larger, with a
much thicker tail and lacking the light neck-stripe.
3. SCIURUS LEUCOPUS FUMIGATUS, subsp. n.
Similar to S. leucopus, but darker, and the outer sides of the
limbs concolorous with the back.
4. FUNAMBULUS RUFIGENIS FUSCUS, subsp. n.
Similar to /. r. typicus, but much darker in general coloration
and having a rufous tinge on the outer sides of the thighs.
A paper was read from Dr. Emin A. Gorxp1, C.M.Z.S., contain-
ing descriptions of seven new or little-known species of Marmoset
Monkeys from the Amazonian Region.
Mr. F. E. Bepparp, F.R.S., read a paper entitled “ Contribu-
tions to the Knowledge of the Systematic Arrangement and
Anatomy of certain Genera and Species of Squamata.”
A communication was read from Mr. Groree H. Kenricsg,
F.Z.S., containing a list, with descriptions of the new species, of
Pyralide collected by Mr. A. E. Pratt in British New Guinea in
1902-03.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 5th February, 1907, at half-past Hight
o'clock p.m., when the following communications will be made :—
1. Prof. E. Ray Lanxsster, F.R.S.—On the Feetus of the
Giraffe.
2. Dr. W. T. Caiman, F.Z.8.—On new or rare Cumacea from
the Collection of the Copenhagen Museum. Part I.
3. Dr. HE, A. Goetp1, C.M.Z.8.—Description of a new Amazonian
Tree-Frog with peculiar Breeding-habits.
The following Paper has been received :—
Mr. C. J. Wirn.—An Account of the South-American Cheli-
Jerine in the Collections of the British and Copenhagen Museums.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society oF Lonpon should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 Hanover Square, Lonpon, W.
January 22, 1907.
No. 39.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON®
February 5th, 1907.
H.G. Tue DuxKE or Beprorp, K.G., President, in the Chair.
Mr. F. Martin Duncan, by permission of the Charles Urban
Trading Co., Ltd., gave a cinematograph exhibition of animals in
the Society’s Gardens and other Zoological Studies, chiefly on the
life-history of Insects.
Mr. Ouprietp Tuomas, F.R.S., exhibited a collection of
Mammals and Birds from the Islands of Saghalien and Hokkaido,
N. Japan, made by Mr. Malcolm P. Anderson in carrying out the
Duke of Bedford’s Exploration of Eastern Asia. Mr. Thomas
proposed to give a full account of the Mammals on a later
occasion.
Mr. Oupriztp Tuomas also read a paper on Mammals collected
in Mindanao, Philippines, by Mr. M. P. Anderson for the Duke
of Bedford’s Exploration of Eastern Asia. Seven species were
mentioned, of which the following was new :—
CRUNOMYS MELANIUS, Sp. n.
Rather larger than C. fallax, of Luzon. Colour wholly dark
blackish brown, not lighter below; limbs and tail also uniformly
dark.
Dimensions of type:—Head and body 98 mm.; tail 68; hind
foot 25. Upper molar series 4:1.
Hab. Mt. Apo, Mindanao. Type. Male. No. 751.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Sic
Shillings per annum, payable in advance.
6
Prof. KE. Ray Lanxester, F.R.S., Director of the British Mu-
seum (Natural History), read a paper entitled “The Origin of the
Lateral Horns of the Giraffe in Foetal Life on the Area of the
Parietal Bones.” The author described and showed the exact
relation of the lateral horns in the fetus taken from the Giraffe
which died last spring in the Society’s Gardens. It was demon-
strated that the lateral horn of the Giraffe was exclusively in
origin a part of the fibrous osteogenetic tissue of the parietal bone
of which it was a part, and had no connection whatever with the
frontal.
Thus the statement made by Su: Richard Owen in his account
of a new-born Giraffe, in a paper read before the Society in 1839,
was finally shown to be based on an unfortunate accident. Owen
had cut out the horn-bearing area of the skull and after an interval
of time had reversed the relations of the excised piece of bone,
taking frontal for parietal and parietal for frontal.
The author expressed the opinion that the parietal lateral horn
of the Giraffe could not be considered to be the same morphological
unit as the frontal lateral horn of the Okapi.
Prof. LANKESTER also read a paper on “ Parallel Hair-fringes
and Colour-striping on the Face of Foetal and Adult Giraffes,” in
which he described, illustrated by lantern-slides, a remarkable
eolour-banding or striping of the hairy covering of the face in the
foetal Giraffe, and showed that similar dark and light striping
occurred in a very marked form in adult Giraffes though not in
all individuals.
Ina third paper, ‘On the Existence of Rudimentary Antlers in
the Okapi,” Prof. Lanxester described the polished tip or apex of
the Okapi’s horn which breaks through the integument. He
showed that transverse fissures or incisions were produced one
behind the other in the naked apex, tending to cut off in succession
a series of small bony caps which he regarded as rudimentary
antlers. He expressly refrained from concluding that this forma-
tion of minute antler-caps was to be regarded as genetically
connected with the antler-formation of the Cervide, though such
a connection was possible.
Prof. LANKESTER also exhibited the skull of a sub-adult male
Okapi obtained by Major Powell-Cotton in the Ituri Forest
(Congo), and a similar somewhat younger male skull, obtained by
Capt. Boyd Alexander in the Bahr-el-Ghazal region. Both skulls
were in a very fine state of preservation.
Mr. G. A. Bountencer, F.R.S8., on behalf of Dr. E. A. GorELp1,
G.M.Z.S., exhibited and described a new Amazonian Tree-Frog,
Hyla resinifictrix, closely related to H. venulosa, but distinguished
by fully half-webbed fingers. This frog was remarkable for its
habit of making good-sized basins of resinous substances in hollow
branches of high trees, in which water collects, which served as a
7
nursery for the eggs and larve. The fzog collected the resin
from the bark of certain trees, such as the aromatic “brew-
branco ” (Protiwm heptaphyllum).
Dr. W. T. Catan, F.Z.S., read the first part of a paper on the
Collection of Cumacea in the Copenhagen Museum. Altogether
30 species were dealt with, of which 25 were described as new.
The majority of the specimens were derived from collections made
in New Zealand and the Gulf of Siam by Mr. H. Suter and
Dr. Th. Mortensen respectively.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 19th February, 1907, at half-past Hight
o'clock P.M., when the following communications will be made :—
1. Mr. R. I. Pocock, F.Z.8.—On English Domestic Cats.
2. Dr. C. G. Seniemann, F.Z.S.—On Deaths occurring in the
Society's Gardens during 1906.
3. Mr. J. T. Cunnincuam, F.Z.5.—On a peculiarly Abnormal
Specimen of the Turbot.
4, Baron F. Nopcsa.—Ideas on the Origin of Flight.
The following Papers have been received :—
1. Mr. C. J. Wrrn.—An Account of the South American Cheli-
Jerine in the Collections of the British and Copenhagen Museums
2. Dr. R. Broom, C.M.Z.S.—On the Dental Succession in the
Cape Golden Moles,
3. Mr. F. E. Bepparp, F.R.S.—On the Azygos Veins in the
Mammalia.
4, Mr. F. E. Bepparp, F.R.S.—On Two new Species of the
African Genus Microchetus belonging to the Collection of Oligo-
cheta in the Museum of Christiania.
5. Miss Dorotuea M. A. Bate.—On Elephant Remains from
Crete, with Description of Hlephas ereticus, sp. nov.
6. Mr. Cuaruzs F. Roussever.—Zoological Results of the Third
Tanganyika Expedition, conducted by Dr. W. A. Cunnington,
1904-05. Report on the Polyzoa.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL SocIETY oF Lonpon should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 HANOVER SQUARE, Lonpon, W.
February 12, 1907.
No. 40.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
February 19th, 1907.
Sir Kpmunp G, Lopzr, Bt., Vice-President, in the Chair.
The Secrerary read a report on the additions that had been
made to the Society’s Menagerie during the month of January
1907.
Dr. C. I. Forsyra Magsor, F.Z.S., exhibited remains of a Bear
from the superficial deposits of a cavern in the mountains of
Corsica, where Bears, though now extinct, were formerly nume-
rous, at least up to the sixteenth century. Despite the fact that
no truly fossil Bears were as yet known from Corsica, Dr. Forsyth
Major considered the Corsican Bear to have been autochthonous,
whilst in his opinion the recent Mammals of Corsica (and Sardinia)
had been, almost without exception, introduced by human agency.
In any case they could not be adduced as proofs of a recent con-
nection of those islands with either of the neighbouring continents.
In a paper on English Domestic Cats, Mr. R. I. Pocock
urged that the surest basis for their classification and the most
satisfactory clue to their descent was furnished by the two dis-
tinct patterns found in so-called Tabby Cats. In one type the
pattern consisted of narrow vertical stripes ; in the other of longi-
tudinal or obliquely longitudinal stripes which, on the sides of
the body, tended to assume a spiral or subcircular arrangement
characteristic of the “blotched” Tabby. This distinction was
long ago pointed out by Blyth.
One or the other of these types was to be found in Cats of almost
all breeds, whether “ Persian,” ‘‘ Short-haired,” or “ Manx.” There
appeared to be no intermediate stages between the two. The Cats
of the “striped” type were no doubt descended from the Kuropean
Wild Cat and the North-African Wild Cat; but the origin of
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance.
10
Cats exhibiting the “ blotched” pattern appeared to be unknown.
It was to the Cat of the latter kind that Linneus gave the name
catus, which was therefore no longer available for the European
Wild Cat; this Cat, therefore, must take the name sylvestris.
Dr. C. G. SznieMANN, the Society’s Pathologist, in presenting
his report on the deaths that had occurred among the Mammals
and Birds in the Menagerie during 1906 stated that 356 Mammals
and 283 Birds were submitted to post-mortem examination, and
the results showed—
(i) That tuberculosis occurring in birds in the Gardens was
usually due to infection by the gut.
(ii) The hearts of Rheas, Cassowaries, Ostriches, and some
of the larger Storks kept in the Gardens were often
extremely flabby, and death in these birds was in a
large number of cases due to cardiac failure.
(iii) New growths were rare both in mammals and in birds,
but one case of carcinoma arising in the kidney and
occurring in a Chilian Pintail (Dajila spinicauda) had
been observed, as well as two instances of benign new
growths occurring in birds not inmates of the Gardens.
Mr. J. T. Cunninenam, M.A., F.Z.8., described a peculiarly
abnormal specimen of the Turbot. The specimen was captured
by Miss Olivia Fox, of Falmouth, near Padstow, on the north
coast of Cornwall. It was a young fish, measuring only 4°4 cm.
in length, and a normal specimen of slightly smaller size, taken at
the same time, was completely metamorphosed to the asymmetrical
condition of the adult. In the abnormal specimen the right side
was almost entirely destitute of colour as in the normal condition,
but both eyes were on this white side, instead of being on the left
side as in normal Turbot. On the left side pigment was present
over the whole surface except the head and the anterior part of
the base of the dorsal fin, which were white. The fish was kept
alive in captivity for two months, and was observed to lie always
with its eyes uppermost, so that the upper side was white and the
lower side coloured. The fish showed also another abnormality,
namely, that the base of the dorsal fin projected anteriorly as a
free process above the dorsal eye, a peculiarity which is usually
present in ambicolorate Turbot. As there was some pigment on
the head on the left side, Mr. Cunningham pointed out that the
specimen might be regarded as a Turbot in which a normal body
was united with a head which was reversed, so that the left side
of the head, bearing the eyes and pigment, was joined to the right
side of the body bearing no pigment, and vice versa.
Dr. Baron Francis Norcsa read a communication entitled
“Ideas on the Origin of Flight,” and illustrated his argument
with lantern-slides showing the hind limbs of various genera of
Bats, Pterosaurs, Birds, and Dinosaurs, as well as a reconstruc-
tion of a hypothetical, cursorial primitive bird. The author
Te
stated that from the mechanical point of view a patagium and a
set of flight-feathers were different organs. He pointed out the
osteological analogies between Bats and Pterosaurs on the one
hand, and between Birds and Dinosaurs on the other. He sug-
gested that Bats and Pterosaurs had arisen from leaping, arboreal
forms, whilst Birds had come from a terrestrial, cursorial stock.
Mr. F. KE. Bepparp, F.R.S., read a paper on the Azygos Veins
in the Mammalia.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 5th March, 1907, at half-past Hight
o'clock P.m., when the following communications will be made :—
1. The Hon. Watter Roruscuinp, M.P., F.Z.8.—Descriptions
of some new Species and Subspecies of Antelopes and of a new
Sheep.
2. Miss Dorornea M. A. Barr.—On Elephant Remains from
Crete, with Description of a new Species.
3. Mr. Cuaruses F, Roussever.—Zoological Results of the Third
Tanganyika Expedition, conducted by Dr. W. A. Cunnington,
1904-05. Report on the Polyzoa.
4, Dr. W. A. Cunnineton.—Zoological Results of the Third
Tanganyika Expedition, conducted by Dr. W. A. Cunnington,
1894-05. Report on the Brachyurous Crustacea.
The following Papers have been received :—
1. Mr. C.J. Wirn.—An Account of the South American Cheli-
ferine in the Collections of the British and Copenhagen Museums.
2. Dr. R. Broom, C.M.Z.S.—On the Dental Succession in the
Cape Golden Moles.
3. Mr. F. E. Bepparp, F.R.S.—On Two new Species of the
African Genus Microchetus belonging to the Collection of Oligo-
cheeta in the Museum of Christiania.
4, Mr. T. A. Cowarp, F.Z.S.-—On the Winter Habits of the
Greater Horseshoe and other Cave-haunting Bats.
Communications intended for the Scientific Meetings of the
ZooLoGicaL Society or Lonpon should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 HANovER Square, Lonpon, W.
February 26, 1907.
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Contents (continued).
February 5, 1907 (continued).
Va ' : Page
Dr. W. T. Calman, F.Z.S. Notice of a paper on New or Rare Crustacea of the Order
Cumacea from the Collection of the Copenhagen Museum ....-.......+204-.ss000% 100
1. The Origin of the Lateral Horns of the Giraffe in Foetal Life on the Area of the Parietal
Bones. By E. Ray Lanxesrer, M.A., D.Sc., LL.D., F.R.S., F.Z.8., Director of the
Natural History Departments of the British Museum ........002,..--00-+--0e-- 100
2. Parallel Hair-fringes and Colour-striping on the Face of Fetal and Adult Giraffes.
By E. Ray Lannester, M.A., D.Sc., LL.D., F.R.S., F.Z.8., Director of the British
Mobseun (Natural History): (Plate V.) 2. ecco. str el wierspets & nbalebaye) eiehalg) a elthele Vere ete 115
3. On the Existence of Rudimentary Antlers in the Okapi. By E. Ray Lanxssrer, M.A.,
D.Sc., LL.D., F.RS., F.Z.8., Director of the British Museum (Natural History).
Gialeipes VB Gc VEL NS ge gees bas 5.0 4 siath does siscaig oco'els hc 2S sie ay rep er 126°
4. Description of Hyla resinifictriz Goeldi, a new Amazonian Tree-Frog peculiar for its
Breeding-habits. By Prof. Dr. Emm A. Gorupr, C.M.Z.S., Director of the Para
AVEUT SESE S2o:\cccvs' vie si aiel al ip'ai ='n wa a\sialsfalaiereisin<cisvale/e\sye & « lele:o/m\egeleiphiie: tet uhe a/carete <laaaeaia mens 135
5, The Duke of Bedford’s Zoological Exploration in Eastern Asia.—lIT. On Mammals
obtained by Mr. M. P. Anderson in the Philippine Islands. By O:prrety Tuomas,
PEE Bose ar ols lc) Salelg initials a sip nia ci ejoie w\nisie\n's e's wiv s d.suslelnh ie ula shat el oiler ke tees 140
February 19, 1907. -
The Secretary. Report on the Additions to the Society's Menagerie during the Month of
Uae US, CEBS 6.46 acto Ap POCO EO BEIEEGBRDe ORDO S a6 go bininieUie'a sie’e) jain cial ama ae etage 143
Dr. C. I. Forsyth Major, F.Z.S8. Exhibition of remains of a Bear from a Cavern in
WOPSIEAS azetaiec « < foe) Sees Sot EGA ee CDR Ones a oabtds. Suopeouesaoéancass . - 143
1. On English Domestic Cats. By R. I. Pococx, F.L.S., F.Z.8., Superintendent of the
Zoological Society’s Gardens. (Plates VIII.-X.) ..-............ picket iwietasyorenyaiats 145
2. Report on Deaths occurring in the Society’s Menagerie during 1906. By C. G. Sztiamayn,
M.D, ) We See atholorist to-the Society... cteic <n: sce ve «0 secs elase areas catahsleeer aaa 168
3. On a peculiarly Abnormal Specimen of Turbot. By J. T. Cunninenam, M.A, F.Z.S.
(CREW: GI ERC APC a ie Ce NIE A ERDAS Coinenmitiy dik GOO HME OO AR eke ac 174
4, On the Azygos Veins in the Mammalia. By Frank E. Bepparp, M.A. (Oxon.),
F.R.S., Prosector to the Society ........... Rcletsietaratatel ets duct acarevefats Map uO Bente eeitc a Lenk
. Ideas on the Origin of Flight, By Dr. Baron Fraxcis Norcsa ..- .
or
LIST OF PLATES.
1907, pp. 1-236.
>
Plate ; Page
BES COncOptBECUS DOMUE Svan e's au So's sao baie: csis Sayers enh 2
TD VEVELCCOD US UOINUS e's sia «0's, bicie's ce ein mnlensl eee ene ele 3
Til.
IV } Pyralidx from British New Guinea ...............-00 - 68
V. Head of a Fetal Giraffe (half the natural size) ........-.. 115
VI. Horn-tips of Okapi ..... BAA Aan aaah dole AON ItoS b's 4 a
VII. Young Ossicone or Horn of Okapi .,..........-++.00..-- } eas
VILE.” Blotched Tabby, Felis catus 2... sie ce vee oo eens ee |
PX Striped: Pabby, Hales Conguata i. « cues cose ok mica eeie iets 143
X. Agriotypes of the Striped Tabby ...... Bea) Sead aah J
xt.) Abnormal Younr: Parbobs. 6s tah 5 oss os = gi + chet lseiecle aii me
NOTICE. .
The ‘ Proceedings’ for the year are issued in four parts, forming two volumes.
as follows:—
Papers read in January and February, in June.
i » March and April, in August.
May and June, in October.
” ”
x » November and December, in April.
‘ Proceedings,’ 1906, pp. 759-1052, were published on April 11th, 1907.
The Abstracts of the papers read at the Scientific Meetings in
January and February are contained in this Part.
PROCEEDINGS
OF THE
| GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
(190%.
Paces 237-446.
CONTAINING PAPERS READ IN
MARCH anv APRIL.
AUGUST 1907.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
_ LONDON:
MESSRS. LONGMANS, GREEN, AND CoO.,
; PATERNOSTER-ROW.
| Yas [Price Twelve Shillings. ]
bees é (
March 5, 1907.
The Hon. Walter Rothschild, M.P., F.Z.S. Exhibition of a mounted specimen of a
Grama e re ee is a eek ots Ge ms spe rahe age By aed Ded ie HEI AN deednat D's oll 237,
1. Descriptions of a new Species‘and two new Subspecies of Antelopes and a new Sheep.
By the Hon. Waurnr Roruscaiyp, M.P., Ph.D., PREZ Gere ctene, Cie iSl siete enere ante cneent ene £237
2. On Elephant Remains from Crete, with Description of Hlephas creticus, sp.n. By
Dorornma M. A. Barn. (Plates XIT. & XTIL.) .. ee eee eee eee ees witha Ale game
,9. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A.
Cunnington, 1904-1905.—Report on the Polyzoa, By Cuaruns FE. Rovussexer,
EVR.M.S. . (Plates XTV. & KV.) cece cece ee cere eee cet ees Bie see astanan aa Soilt cet 250°
4. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A
Cunnington, 1904-1905.—Report on the Brachyurous Crustacea. By Winwiam A.”
Cunninoron, B.A., Ph.D., F.Z.S. (Plates XVI. & XVIL.)...- ee eee ee eee ee eee 208
5. On Two new Species of the African Genus Microchetus belonging to the Collection of ,
Oligocheta in the Museum of Christiania. By Frank E, Bepparp, M.A., F.RS,,
Prosector to the Society ........ esse ee ee ee ee ee ee es BA IOS Us NE a eee Ea QT
March 19, 1907.
The Secretary. Report on the Additions to the Society's Menagerie during the month of
February 1907.2... ce cece cee ee tener eens Bea chat obs calanecalcale fe] cuss catetote 281
-Mr. Herbert F. Standing. Notice of ‘a Memoir on recently discovered Subfoussil Prosimiz
from Madagascar ........ SEN APT ER NADL OR EAL gisem cunts AAG et --- 28h
1. Descriptions of some New Species of Animal Parasites. By L. W. Samson, M.D., HAS. 282
2. Descriptions of five New Species of Hamogregarines from Snakes. By L. W. Sampon,
M.D., F.Z.S., and 0, G. Seniamann, M.D., F.Z.8. ........ 0 Ph Reape ec Sees 28
3. The Rudd Explor ation of South Afvica,—VII. List of Mammals obtained by Mr. Grant
at Coguno, Inhambane. By Oxpriztp Tuomas, F.R.S., F.Z, S,,and R. C. Wrovenron,
1S AIS Ce er a @eneese ae beapeeeeereenere ee een enews ee ey 285
Contents pomied on page 3 of Wrapper.
1907. | ON NEW ANTELOPES AND SHEEP. 23
March 5, 1907.
Freperick Giniert, Ksy., Vice-President, in the Chair.
The Hon. Walter Rothschild, M.P., F.Z.S., exhibited a mounted
specimen of a Gorilla, Gorilla gorilla diehli.
The following papers were read :—
1. Descriptions of a new Species and two new Subspecies of
Antelopes and a new Sheep. By the Hon. Waurer
RorascHiLp, M.P., Ph.D., F.Z:8.
[Received March 3, 1907.}
RHAPHICEROS HORSTOCKI NATALENSIS, subsp. n.
Differs from 2. horstocki in its much darker colour. The white
patches on the throat and round the eyes much smalier and the
white on the belly less extended. The whole of the rest of body,
head, and limbs dark vinaceous rufous instead of orange-rufous.
Hab. Neighbourhood of Drakensberg, Natal.
CERVICAPRA ARUNDINEUM OCCIDENTALIS, subsp. n.
Differs from C. arundinewm in the paler, more greyish rufous
of head and neck and the pale rusty-grey of limbs, tail, and body.
The horns also seem straighter and thicker as a rule.
Hab. Neighbourhood of Fort Jameson, Northern Rhodesia, and
Bangweolo Flats.
CoBuUs ROBERTSI, sp. 0.
IT am treating this as a species because it occurs in the same
territory as true C. lechwe and C. smithemanni, otherwise it has
many characters intermediate between these two species. Nor
do I consider it likely that it is a hybrid, for it has been shot by
several sportsmen, and not always in the same localities.
Horns stouter and the rings broader and closer together than
in ©. lechwe, smooth tip shorter. Head, hind-neck, body, and
upper part of fore-neck bright orange-rufous, back and flanks
darker and redder than in C. lechwe. Cheeks, throat, and sides
of neck have the rufous strongly mixed with black hairs, thus
somewhat approaching C. smithemanmi. Lower fore-neck whitish,
with two large black patches on each side, which join similar
patches on the front of the shoulders. Outside of ears rafous
buff, inner side white.
Hab. Northern Rhodesia. (Type, collected by Mr. Roberts.)
Proc. Zoou. Soc.—1907, No. XVII. AF
238 MISS D. M. A. BATE ON [ Mar. 5,
OvIs COWANI, sp. n.
This Sheep is nearest to O. stonrei, but differs in being entirely
deep black, with the exception of a white rump-patch and a grey
face. The rump-patch is considerably smaller than in O. stonet.
The three-year old ram of O. stonei is occasionally much darker
than a specimen of that species in my possession, but is always
of a rusty or brownish black, and has the very large white rump-
patch.
The type specimen of O. cowani was entire, but the skin had
only been dried and not dressed, and when relaxed the whole of
the hair came off, and only the head and neck could be saved.
The type, shot by an Indian out of a large flock, was sent
me by Mr. C. G. Cowan, of Kamloops, and was obtained in the
mountain-chain near Mount Logan in British Columbia.
Types of all these four species and subspecies are in the Tring
Museum.
2. On Elephant Remains from Crete, with Description of
Elephas creticus, sp.n. By Dorornea M. A. Bats.*
[Received February 1, 1907. |
(Plates XII. & XIII.7, and.Text-figure 83.)
Introduction.
Perhaps the most important and interesting of the results of
the author’s visit. to Crete in 1904 was the discovery, in two
hitherto unexplored cave-deposits, of the remains of Elephants of
different sizes, more particularly as the occurrence of one of these,
of pigmy proportions, appears not to have been previously known.
Although prior to 1904 no large quantity of specimens seems
to have been obtained, yet the existence of ossiferous deposits
in this island has been known for a considerable time, as the
following records testify. The earliest would seem to be that of
Pococke, who described a bone-cave in the Khania Akrotiri in a
volume published in the middle of the eighteenth century 2.
Nearly a hundred years after this, a reference occurs relating to
fossilised human remains found, together with marine forms, near
Khania by Fabrequette §, who was at one time Consul at Malta.
Later, remains of a Hippopotamus, which prokably came from
the upland basin of Lassethe, were obtained by more than one
traveller ||, and have since been referred to by a number of writers.
Two ossiferous caves were discovered in the west of the island by
* Communicated by Henry Woopwarp, LL.D., F.R.S., V.P.Z.S., F.GS.
+ For explanation of the Plates, see p. 250.
+ Richard Pococke, ‘A Description of the East’ (London, 1745), vol. 11. p. 264.
§ C. R. Acad. Sci. (Paris), iv. 1837, p. 182; also ibid. viii. 1839, p. 178.
|| See Admiral Spratt. ‘Travels aud Researches im Crete’ (London, 1865), vol. 11.
pp- 386-7; also Raulin, ‘ Description Physique de l’Ile de Créte’ (Paris, 1869), vol. i.
p. 156, and vol. 11. p. 615.
Das, WOW, IPL, 20.
SIOZCH
nat
West,Newman imp.
G.M-Woodward del.et lith.
o
FLEPHANT REMAINS FROM CRETE.
EhHaO WOU SNIVNEY GUNVHde Ta
-durr weurms yy “4Sse “UQIT 99" TOpP Puempoom WD
“TINDSC Weal, AA OWS SIE TS) 74 val
1907. ] ELEPHANT REMAINS FROM CRETE. 239
Admiral Spratt *, both of which have since been visited by the
present writer ‘.
Signor Simonelli, whose geological researches in Crete were
carried on in 1893, was seemingly the first to obtain Elephant
remains, which he procured from cayes near Retymno, which is
situated on the north coast and les between Khania and Candia.
In a paper published in 1894 = he identifies the specimens, which
included a perfect mandibular ramus, as those of Elephas priscus
(=4. antiquus §), though no detailed descriptions or figures are
given, and I have been able to find no other published reference
to them.
Unfortunately the collection of teeth and limb-bones which
forms the subject of the present paper consists of a small amount
of material only, many of the specimens being but imperfectly
preserved. Notwithstanding this they fall naturally into two
groups representing animals of different sizes, the smaller of
which agrees in this respect with H. melitensis, whilst a number
of limb-bones indicate an Elephant superior in dimensions to the
average Indian species (#7. maximus) of the present day, but not
equalling the gigantic proportions at times attained by Z. meri-
dionalis and L. antiquus, though, as will be shown later, it is to
this last that they must evidently be referred. Thus at first sight
it seemed probable that the collection included remains of both a
dwarted race and the parent form from which it had sprung.
However, before remarking on the relationships suggested by a
study of the specimens obtained, it will perhaps be more con-
venient first to briefly describe them, commencing with those by
which the pigmy form is represented.
I. ELEPHAS CRETICUS, sp. n.
The remains of the smallest of the Cretan Elephants were all
obtained from a much damaged and weathered cave-deposit in the
limestone cliffs near Cape Maleka in the west of the island, which
has already been described ||, and where only some teeth and
limb-bones of small rodents were found besides those under dis-
cussion. These latter include nine imperfect molars and a few
fragments, among which are a portion of an incisor and the
dorsal half of a vertebra. As this small race differs from those
of other Mediterranean islands, and its minute proportions being
seemingly the result of specialisation due to isolation in Crete, it
is suggested that it may be known by the above specific name
denoting its island habitat.
Incisors—Of the milk-incisors no specimen was _ procured,
while, as already remarked, only a fragment of the proximal end
of a permanent tusk (M. 9375) was obtained. From the limited
* Op. cit. vol. u. pp. 194-5.
+ Geol. Mag. n. s. dec. v. vol. 11. (1905) pp. 194-6.
t Atti R. Accad. Lincei, 5* ser. Rendiconti, vol. i. 1894, Sem. 2, pp. 265, 268 ;
also ‘Candia’ (Parma, 1897), pp. 171-2.
§ Cat. Foss. Mamm. Brit. Mus. part iv. (London, 1886) p. 122.
|| Geol. Mag. n. s. dee. v. vol. 11, (1905) p. 195.
ys
240 MISS D. M. A. BATE ON [ Mar. 5
extent of the pulp-cavity, it is believed to have belonged to a nearly
adult individual and indicates a tooth of small size, though its
condition is too imperfect to admit of any measurements being
given other than its approximate diameter, which is 46 mm.
Upper Molars. ——(Ornlhy two of the nine sanvellanis are considered to
belong to the upper series; the smaller of these (M. 9376), a first
true molar, i is split longitudinally aimost in half, but sufficient
remains to show the general character of the tooth, It consists
of seven plates, though it is possible that an additional small
anterior ridge was originally Breseut: Its antero-posterior length
is 54 mm, and height about 32 mm., although the fangs s extended
for a short distance further, bringing the total measurement up
to 41 mm, Comparatively speaking, the height of this tooth is
slightly greater than in most of the other specimens referred to
E. creticus; this is an argument in favour of its being a first true
molar rather than the last of the milk- series, which, from its size
alone, it might perhaps have been thought to represent. The
enamel bands are not very thick and im several of the ridges are
somewhat broken up into “rings”; at the same time, the cement-
areas are broad, a character generally found in conjunction with
more massive enamel. In section this tooth shows that the cement-
areas maintain an almost uniform width for nearly the whole of
their extent. This feature is probably not infrequent in the
molars of this species, as suggestively exemplified by some of the
extensively worn specimens described below.
The Cretan specimen is superior in antero-posterior length to an
example of the corresponding tooth of H. cypriotes (M. 8602) *,
though this latter exceeds it in the height of its crown, which
is 53-5 mm.; this difference in the proportions is even more
marked in a molar of 1, melitensis (44252) r.
The second specimen (M. 9577, PI. XCUL, fig. 2), believed to
belong to the upper series, is of larger propor tions than the last,
and is much worn and somewhat damaged ; it probably represents
the penultimate true moiar. tis difficult to say of how many plates
it originally consisted, but it can be seen that there were at least
eight, all of which were in use, the front ones bemg worn down
almost to their common base, while a portion of the anterior ridge
is broken off. Partly owing to its advanced state of wear, the
crown is very wide, being 37 mm. across, while its antero-posterior
length is 77 min. and its height 37 mm., though this would
of course have been greater when the tooth was still but slightly
abraded. In spite of the condition of this molar the cement-areas
are still broad, equalling or even exceeding in size the plates of
dentine; the enamel bands are thick and in a few instances
somewhat wavy in outline.
Aithough apparently considerably superior in height of crown,
this tooth in 4. cypriotes is otherwise decidedly smaller than the
above, the dimensions of a much less worn example (M. 8601)
* Fioured, Phil. Trans. vol. 197 B (1904), p. 351, pl. 21. figs. 4, 4a.
+ Trans. Zool. Soc. vol. ix. p. 20, pl. 2. figs. 9, 9a.
1907. ] ELEPHANT REMAINS FROM CRETE. 241
being :—approximate antero-posterior length 72 mm., greatest
width of crown 28 mm., and total height 65 mm. Compared
with the specimen (49267) * identified by Falconer as M. 1 of
LE. melitensis, this last is somewhat narrower in the crown, which
is 53 mm. across, though this may be partly accounted for by the
difference in age, and it is likely that the two would originally
have closely agreed in this respect. On the other hand, in height
the molar from Malta greatly exceeds the one from Crete, being
very high in relation to its width, as already noted by Falconer ;
further, the cement-areas in the Cretan fossil are distinctly
broader.
Lower Molars.-—Ot the seven molars obtained not one can be
referred to a position among the milk-teeth, while, owing to
imperfect preservation, it is impossible in one or two cases to make
certain of the place originally occupied in the permanent series.
The first true molar appears not to be represented. Two specimens,
both belonging to the right side of the mandible, are believed
to be second true molars. One (M. 9578 a) has been considerably
weathered and is much worn, all except the last of the plates
having been in use; the number of these was at least nine. The
‘cement-areas are broad and the enamel bands thick and smooth,
while in two, if not three, of them there is a median loop,
though this is unaccompanied by a mesial expansion of the plates
of dentine.
The other tooth (M. 9378, Pl. XIIT. figs. 1, 1 a), regarded as a
second true molar, is also extensively worn, the penultimate ridge
being already slightly abraded ; although it is broken off anteriorly
seven plates remain as well as a portion of an eighth. The width
of the seventh plate (counting from the rear) is 37 mm., while the
height of the preantepenultimate one is only 33 mm. : the enamel
is massive and in the four posterior used plates is somewhat
irregular, while the cement separating the anterior ridges is con-
siderably excavated. A comparison shows this Cretan specimen
to be broader and much lower in the crown than a corresponding,
though less worn, tooth of £. eypriotes (M. 8588) 7
Judging from their size it is probable that two other lower true
molars, very imperfectly preserved, also represent the second of
the permanent series. One of these (M. 9382) is situated in the
posterior half of a left mandibular ramus; it probably consisted
of eight or nine plates and, like most of the teeth obtained of
E. creticus, is low-crowned and has thick enamel bands and wide
cement-areas. The other example (M. 9380) is a portion of a much
damaged right molar; the worn surfaces of two ridges remain and
seem in no way to differ from those of the specimens already
described.
The last lower molar (M. 3) is represented by three specimens,
none of which is in an advanced stage of wear. Two of these,
* Falconer, Pal. Mem. vol. ii. pl. 11. figs. 2, 2a, p. 294; also identified by Busk as
M. 1? of #. falconeri, Trans. Zool. Soc. vol. vi. part v. pl. 53. figs. 9, 9 a.
+ Phil. Trans. vol. 197 B (1904), p. 355, pl. 21. figs. 3, 3a.
242 MISS D. M. A. BATE ON [Mar. 5,
damaged anteriorly, are each situated in a portion of a ramus;
and belonging, as they do, to opposite sides of the mandible as well
as being in a similar stage of wear, it is thought that they may
have been owned by a single individual. On the left side
(M. 9383, Pl. XII. fig. 1), in addition to the last molar, the
three posterior plates of M. 2 are also present. In M. 3 eleven
plates can be counted, but a considerable part of the posterior
portion of the tooth is wanting. The fifth plate is slightly abraded,
and it is noticeable in both this and the companion specimen that
the enveloping cement on the grinding-surface of the crown is
somewhat scanty, with the result that all the enamel soon becomes.
exposed, even while the anterior portion of the tooth may still be
but little worn. The second example (M. 9383 6) presents very
similar characteristics ; the presence of eleven plates can be deter-
mined, but damage anteriorly and concealment of the tooth behind
py the boneand matrix make it impossible to ascertain the original
total.
The one other last true molar (M. 9381, Pl. XII. fig. 3), an
isolated tooth of the left side, is almost perfectly preserved except
for the loss of the greater part of the first and a portion of the
second plate. It is considerably curved, this being accentuated by
the angle at which the hinder plates lie. The number of ridges is
thirteen, six of which show signs of more or less wear, and there
appears to be no evidence of the former presence of any additional
ones. The sloping, instead of upright, position of the last plate
is a further indication that this molar must be considered an
example of the last of the permanent series. The antero-posterior
length of the tooth (not along the curve) is about 122 mm., or
139 mm. if continued to the heel of its base; the greatest height,
which occurs at the sixth plate, is 53 mm., while the width of the
abraded surface of the third ridge is 3) mm. The cement-areas
are of medium width and the enamel bands thick and uncrimped,
though somewhat irregular and disconnected in the less-worn
plates.
It is interesting to contrast this with two corresponding teeth
of #. cypriotes in the British Museum Collection *, ail three being
of the left side and in practically a similar stage of use. The
Cretan fossil is the more massive and superior in width and
antero-posterior length, the persistence of the annulation of the
enamel bands is also more strongly marked. On the other hand,
it has a lower crown (53 mm.), both actually and as compared
with its bulk, than the Cypriote specimens, this being 59 mm.
in. the isolated tooth (M. 8591), while in the other (M. 8589),
although the ramus prevents a measurement of the greatest
height being obtained, that of the seventh plate is 63 mm.
The most satisfactorily identified last molar ef H. melitensis in the
British Museum Collection for comparison with the above-noticed
tooth from Crete, is a specimen situated in a right mandibular
* M. 8589, Phil. Trans. vol. 197 B (1904), p. 355, text-figs. 2&3; and M. 8591,
ibid. p. 355, pl. 22. figs. 6, 6a.
1907.] ELEPHANT REMAINS FROM CRETE. 243:
ramus (M. 44294), already described and figured by Dr. Leith
Adams*. It is evident that in antero-posterior length it
slightly exceeds the Cretan example and consists of several more
plates, these being in much closer proximity owing to the narrow-
ness of the cement-areas and the less massive enamel bands.
Considering the size of the ramus this tooth must be much the
higher of the two, while in breadth of crown the molar from Crete
is superior,
The one other dental specimen obtained (M. 9379, Pl. XII.
fig. 2) consists of only three unworn plates; the anterior surface
of the foremost of these is strongly but simply grooved, its greatest
width is 35 mm. and its height 50 mm.
The only bone procured from the deposit near Cape Maleka 1s
the dorsal half of a vertebra embedded in the matrix attached to
the mandibular ramus containing a last true molar (M. 9383).
From this brief account of the remains procured of H. creticus,
it will be seen that this pigmy Elephant must have been of slightly
larger proportions than #. cypriotes and approached in size more
closely to H. melitensis ; that is to say, it would have attained as
a maximum a height of five feet f. All the molars obtained differ
from those of the two last-named dwarf species in being much
lower in the crown; this is perhaps the most noticeable feature of
the series. At the same time the teeth are wide, the cement-areas
broad, and the enamel simple, though at times broken up into a
number of rings. So far as can be ascertained from the scanty
amount of material the ridge-formula must have been low.
Except with regard to the immense difference in size the
characteristics of the molars described above, more especially in
the lowness of the crowns ¢, appear to resemble more closely those
of H. meridionalis than of any other of the larger Elephants of
the Mediterranean Region.
I]. Eneruas antiguus Falconer.
As previously mentioned, remains of this Elephant $ had already
been obtained from caves closeto Retymno. The teeth and bones
noted below, and believed to belong to this species, were all pro-
cured from a much damaged and fragmentary cave-deposit, one
of several found close together in the cliffs bordering the south
of Kharoumes Bay in the Eparkhia of Sitea|. Although evi-
dently but a remnant of a formerly larger deposit, it was possible
* Trans. Zool. Soc. vol. ix. p. 30, pl. 6. figs. 1, la.
+ Leith Adams, Trans. Zool. Soc. vol. ix. pp. 108, 116.
{ This characteristic of the molars of H. meridionalis was constantly noted by
Falconer, see Pal. Mem. (London, 1868), vol. ii. pp. 128, 134, 138, &c.
§ Signor Simonelli does not give the author of his H. priscus, but it may be sup-
posed that this name is employed for the thick-ridged variety of EH. antiquus
(Falconer’s HZ. priscus, Cat. Foss. Mamm. Brit. Mus. part iv. p. 122), since the name
E. priscus of Goldfuss appears to have been applied to molars almost indistinguish-
able from those of H. africanus (see Pomel, Bull. Soc. Géol. France, tome vii.
1878, p. 51).
|| Geol. Mag. n. s. dec. y. vol. i1. (1905), footnote 3, p. 199.
244 MISS D. M. A. BATE ON [ Mar. 5,
to trace the presence of Hlephant bones for a depth of several feet.
Some few remains of ruminants, similar to those found in other
parts of the island, also occurred here, but were only observed
close to the uppermost of the bones of the Proboscidian, which
probably became extinct long before these smaller mammals.
Jt has already been mentioned elsewhere * that in another cave-
deposit, on the same level—that is to say, not many feet above
the sea, and only a few yards distant from the one under dis-
cussion,—were found a number of land-shells, Helix pellita Fer., a
species seemingly not previously recorded in a fossil state 7, these
being preserved in the hard red breccia so common in cave-deposits.
The presence of these shells points conclusively to the deposition
of the mammalian remains under land-conditions, though it now
appears just possible that these deposits were subsequently at all
events partially submerged, which may help to account for their
fragmentary condition. The occurrence of this movement is
suggested by the discovery, in some sand adhering to a femur of
HE. antiquus, of a large number of foraminifera and other marine
forms, an account and list of which have been published by the
Rev. R. Ashington Bullen=. Traces of a former submergence are
more noticeable in the west of the island, especially at Sphinari
and the Kutri and Haghios Basilis caves ; it was also unexpected
in the east, where the coastal movement is supposed to have been
for long past in an opposite direction to that in the west §.
H. antiquus is represented by a number of limb-bones, including
several perfectly preserved foot-bones, and a single right man-
dibular ramus (M. 9384) containing twosomewhat damaged molars.
Both from its size and from the fact of its having been found
situated just above the limb-bones, it was at first sight thought
that this last indicated the former occurrence in Crete of a small
race of Elephant intermediate in size between the pigmy /. creticus
and the very large species indicated by the remains noticed below.
However, a further study of the material has caused the conclusion
to be reached that it is a portion of the mandibular ramus of an
immature specimen of #. antiguas, and that the teeth must be the
penultimate and ultimate milk-molars, or perhaps the last of the
milk and first of the permanent series. The general appearance
and characters of these molars support this view of their identity,
which is further strengthened by the fact of the large limb-bones
occurring in the same deposit, and also that H. antiqwus has already
been recorded from another district of the island.
Lower Molars.—As already remarked, the two lower molars
obtained, and believed to be those of #. antiquus, are situated in
a portion of the right mandibular ramus shown in text-fig. 83
and Pl. XIII. fig. 3: neither of these teeth is quite complete.
* Geol. Mag. n. s. dec. v. vol. ii. 1905, footnote 2, p. 199; and Rev. R. Ashington
Builen, Proc. Malacol. Soc. vol. vi., Sept. 1905, p. 307.
+ Ibid.
£ Geol. Mag. n. s. dec. v. vol. iii. pp. 353-358, pls. 18 & 19.
§ For references to this, see Geol. Mag. n. s. dec. v. vol. ii. (1905), footnotes 2 & 3,
10s IZ,
ES Orie ELEPHANT REMAINS FROM CRETE. 245
The whole specimen is about 263 mm. (nearly 103 inches) in
length ; the ramus is robust, the greatest thickness in the portion
preserved being 112 mm., and its depth, measured in front of the
first molar, is about 126 mm. Its anterior border is very abrupt
owing to the advanced state of wear and consequent forward
position of the anterior of the two teeth.
Unfortunately neither of these molars retains its full com-
plement of plates, which makes it impossible to determine with
absolute certainty their exact position in the series, for, as may be
seen by the specimens in the British Museum Collection, and has
been pointed out by Dr. Leith Adams*, the milk-teeth of this
species varied in size to a very great extent. However, it seems
certain that the specimens in question represent either the two
last milk-molars or the posterior of these and the first true molar.
Text-fig. 83.
Right mandibular ramus of Hlephas antiquus (2) bearing two lower molars.
1 = .
3 hat. size?
3
The actual height of the foremost tooth (Pl. XIII. fig. 3 and
text-fig. 83) is not shown, owing to the enveloping ramus; it
consists of eight plates, but is extensively worn and _ projects
considerably beyond the edge of the alveolus, so it is likely thate
there may have been one, perhaps two, additional plates originally.
The present length of the crown, measuring along the median
line of the plates, is 92 mm., and its greatest width, which occurs
at the sixth of the plates present, is 47 mm., its height above the
ramus at the same place is 30 mm. ‘The cement-areas are not so
wide as those of the following molar, but this is probably due to
difference in‘vear ; in neither is there any mesial expansion of the
plates. The enamel bands are rather thin and “‘ wavy” in outline,
in both these respects differing from the specimens from Cape
Maleka.
The second of the two teeth also shows eight plates, though it
* Mon. Brit. Fossil Elephants, Pal. Soc. London, 1877.
246 MISS D, M. A. BATE ON [ Mar. 5,
is evident that one or more may be wanting posteriorly. Only the
anterior five are worn, and the enamel is seen to be less irregular
than in the preceding molar, considerable digitation of the plates
is observable, while the cement-areas are of considerable width.
This molar is remarkable for its very great height, being 103 mm.,
while the greatest width of the crown at the second plate is only
43 mm. It will be remembered that this was given by Falconer *
as one of the distinguishing characteristics of the molars of
E. antiquus :—*‘ Great height of the plates. The height is more
than double the width of the crown.” This is in striking contrast
to the proportions of the teeth of Z. creticus, in some examples of
which the height of the crown is exceeded by its width. Particu-
larly in the rather thin and very wavy outline of the enamel bands,
these specimens from the Kharoumes deposit resemble a number
of those of H. mnaidriensis figured by Dr. Leith Adams 7, who
also called attention to the resemblance between the molars of this
Maltese species and those of 1. antiguas £.
Limb-bones.—The specific or peculiar characters of the limk
bones of #. antiquus do not appear to be well known, partly no
doubt owing to the difticulty of distinguishing them in cases where
the remains of more than one species of similar size occur in a
single deposit, so that it is evidently chiefly by inference that the
fragmentary collection of bones under discussion must be deter-
mined as those of this species. However, it will have to be
acknowledged that this contention is a strong one when we con-
sider the identity of the teeth found in the same deposit, and the
discovery of the cave near Retymno from which were obtained
similar, though more complete, remains, At least two individuals
are represented amongst the limb-bones, which number about
twenty and are almost all imperfect, with the exception of a few
foot-bones ; and in many cases the articnlar surfaces are damaged
or missing, which makes it difficult to discern any features other
than that of size. Nearly every specimen was covered with a thin
red stalagmitic encrustation.
The only portion of the spinal column procured is a portion of
the neural arch of a dorsal vertebra (M. 9388). The collection
contains two wlae (M. 9385), though only the proximal portion of
each is preserved. One of these, which is unfortunately much
crushed, is of the right side and apparently that of an adult. It
seems to agree in size and general appearance with a corresponding
bone of #. antiquus in the British Museum Collection (45203) §.
A fragment of the humerus is still attached to the second specimen,
which belongs to the left side and is that of a young individual,
the line of junction between the shaft and the olecranon epiphysis
being very apparent. On comparing it with the proximal portion
of a left ulna in the Collection of the British Museum (45202) |j,
* Pal. Mem. vol. ii. p. 176.
+ Trans. Zool. Soc. vol. 1x.
+ Mon. Brit. Fossil Elephants, Pal. Soc. 1877, pp. 25 & 50.
§ Ibid. p. 59, D. 12. || Lbid.
HO Oral ELEPHANT REMAINS FROM CRETE. 247
it appears that the depression below and between the olecranon
and the projection of the sigmoid notch is more extensive in the
former, and also that the distance between the coronoid process
and the olecranon is comparatively greater.
An imperfectly preserved left femur, with its distal extremity
missing, was obtained, also the head of another and a fragment of
the head of a third. The entire length of the former is 29 inches
(89 mm.); it is decidedly short in the neck and has a shallow
digital pit, both of which characters are claimed for #. antiquus by
Dr. Leith Adams*, as well as for ZH. namadicus t+, E. africanus,
and #. mnaidriensis.
A left cuneiform, which is attached to the unciform, is very
much larger than a corresponding bone in the British Museum
Collection (36608) +, which, however, is by no means a large
example for this species. The greatest width of the Cretan
specimen, from its inner border to the outer angle, is 165 mm.,
while antero-posteriorly it measures 115 mm., though this would
be slightly more were it quite intact. The height of its
anterior face is 57 mm. The greatest height of the anterior
face of the above-mentioned wnciform is 81 mm., antero-
posteriorly it is 125 mm., and from side to side 151 mm.
A considerably damaged right unciform appears to be of similar
proportions.
An almost perfect right trapeziwm included in the collection
is found to differ considerably in size and outline from the one
figured by Dr. Leith Adams, and considered by him to present
diagnostic characters §. The former is altogether much smaller
and slighter than the latter, its greatest height, taken perpen-
dicularly, being 79 mm., and its circumference just below the
external articular surface 180 mm. In the British Museum
specimen the lower articular surface || is expanded and almost oval
in shape, whilst in the one from Crete this is narrower and some-
what elongated. The projection beyond the anterior border of the
distal articular surface is extremely pronounced, and it is chiefly
to this that the bone owes its irregular outline.
The metacarpals ave represented by two distal extremities
and an almost perfect specimen of the fifth metacarpal of the lett
manus. Its exterior border is 142 mm., the proximal articular
surfaces measure 83 mm. from inner to outer edge, while antero-
posteriorly they are about 78 mm., the circumference at the centre
of the shaft is approximately 210 mm.
The proximal phalanx of the fifth digit of the right manus,
another digital phalanx, an imperfect right semilunar, and some
portions of ribs, are the only other well-preserved remains of
EL. antiquus obtained.
* Mon. British Fossil Elephants, p. 63.
+ Considered by some writers as a variety of or identical with EH. antiquus (see
G. E. Pilgrim, Records Geol. Survey India, vol. xxxii. part 3, pp. 204, 205, 215, &e.).
+ Cat. Foss. Mamm, Brit. Mus. part iv. p. 136. ;
Ab
§ Op. cit. pp. 160 & 234, pl. 19. figs. 9-9 B, B.M. 20821.
|| Led. fig. 9B.
248 MISS D. M. A. BATE ON [ Mar. 5,
TIL. ConcLustons.
It was at one time thought that the discovery of remains of
different Elephants in the Pleistocene cave- deposits of Crete might
help to throw additional, or perhaps even fresh, light on the subject
of the origin and development of pigmy forms. However, it will
have been seen from the above descriptions that the material ob-
tained is very fragmentary, so that little more than the identity
of the species can be established, though there is at least sufficient
to suggest one or two minor problems of some considerable interest.
The occurrence of two species of Elephant in more or less con-
temporary deposits in such a comparatively restricted area as Crete
hints at, though by no means proves, the likelihood of some
possible relationship or connection between them. It seems quite
certain that Crete, evenif well covered with a luxuriant vegetation,
could not support for long an Elephant of such great size as
Fi. antiquus, so it 1s probable that this large race, at all events as
such, ceased to exist in early Pleistocene times, which would be
tantamount to saying shortly after the separation of the island
from the mainland.
It seems a firmly established and well-supported theory that
the dwarf proportions of the pigmy species found in most of the
larger Mediterranean islands are a specialised character acquired to
meet the exigencies of these island habitats ; for, as M. Gaudry has
remarked *, these former small races have confirmed the idea that
the size of animals was in accordance with the extent of their
habitat. Among other arguments, if needed, in favour of this is the
fact that up to the present no similar remains of a small form are
known to occur in deposits earlier than Pleistocene. In this con-
nection further information with regard to the extent and cause
of the dwarfing of H. africanus pumilio Noack 7, from the French
Congo, will be of the utmost interest, and might also help to
elucidate the problem of the possibly analogous Hippopotamus
liberiensis.
Nowadays many writers maintain that #. antiqus is the parent
form of the hitherto described pigmies of the Mediterranean aren,
and this seems to have been well established by Prof. Pohlig = in
the case of the remains from Sicily. At first sight the idea that
these tiny forms were all derived from one of the largest species
of the genus appears to be a somewhat startling one, but may it
not be that the attainment of such dimensions under favourable cir-
cumstances points to the species having been one highly susceptible
to and strongly acted upon by its environment and conditions of
life? If so, the same would apply under opposite ee
adverse circumstances causing a rapid diminution in size; for, a
* “Ont confirmé Vidée que la taille des animaux était en rapport avec l’étendue
de leur habitat” ‘Foss. de Patagonie, Ann. de Paléontologie, tome i. fase. 11., Juin
1906, p. 8.
+ Zool. Anzeiger, Bd. xxix. No. 20, Jan. 1906.
ft Abh. d. k.-bay. Akad. Wiss. xviii., and Nov. Act. L.-C. Akad. Naturforsch. lvii.
POO ELEPHANT REMAINS FROM CRETE. 249
Mr. Pilgrim has remarked*, it is to this adaptability that
E. antiquus owed its wide geographical distribution and also its
continued existence through a long period of time.
Therefore, allowing that ZH. creticus is a dwarfed species, the
question of descent is decidedly the most interesting point raised
by this newly discovered pigmy of Crete. Two alternative theories
with regard to this subject suggest themselves :-—
A. That, taking into consideration the several points of resem-
blance, which may prove to be only superficial, between the
molars of ZL. creticus and HL. meridionalis, the latter may be
the parent form, in which case it must have inhabited Crete
contemporaneously with #. antiquus.
B. That the Cretan race is descended from Z. antiquus, which
we think is most likely to prove the true solution of the
question. Some few points which appear to support this
may be noted here. Among these are the fact of remains
of this large species being found in the island and the
evidence which tends to show a similar ancestry for other
island races of the same region.
Supposing #. creticus to be derived from Z. antiquus, this would
necessarily mean that the former became differentiated since early
Pleistocene times. That this calculation would allow ample time
for such a change to be accomplished seems probable, when it is
considered that /. eypriotes was the result of isolation for a no
longer period, since, according to Messrs. Bellamy and Jukes-
Browne *, it seems that a land-connection existed between Cyprus
and the mainland as late as the beginning of the Pleistocene era.
Both from the geological evidence and from the apparently still
plastic condition of the small Elephants of Malta, as well as of the
Hippopotamus of Sicily, at the time of their extinction, it is likely
that these islands, more particularly the latter, have formed areas
of isolation for an even shorter term than Cyprus. Therefore it
would have been vital to the continued existence of the species
that the dwarfing should be accomplished as rapidly as possible ;
this also applies to the molars, in which the result might have
been attained by diminishing the height and number of the plates,
thus apparently to a certain extent reverting to a more simple
condition. A significant fact which may lend weight to this sug-
gestion is that, although #. mnaidriensis and #. melitensis of
Malta are seemingly closely connected and probably represent two
stages in the evolution of a pigmy from a large Elephant, yet the
latter has the lower ridge-formula =, although it is the smaller and
therefore, in that direction, the more specialised of the two.
My sincere thanks are due to Dr. A, Smith Woodward, F.R.S.,
* Records Geol. Survey India, vol. xxxii. pt. 3, p. 216.
+ ©The Geology of Cyprus, chap. viii. (Plymouth, 1905); and Bellamy, Key to
Geol. Map Cyprus, pp. 15, 16 (Stanford, 1905).
t Lydekker, Cat. Foss. Mamm. Brit. Mus. part iv. p. 151.
250 MR. C. F. ROUSSELET ON THE POLYZOA [ Mar. 5,
for giving me every facility for working in his department of the
British Museum (Nat. Hist.).
EXPLANATION OF THE PLATES.
Puate XII.
Fig. 1. Inner side of the posterior part of the second and anterior part of the third
right lower molars of Hlephas creticus. (M. 9383a.) | nat. size.
2. Unworn plates of a molar of H. creticus. (M. 9379.) Nat. size.
3. Crown view of right lower third molar of 4. ereticus. (M. 9381.) nat.
size.
Prats XIII.
Fig. 1. Crown and (1 a) side views of second lower molar of H. creticus. (M. 9378.)
+ nat. size.
2. Crown view of second upper molar of #. ereticus. One of the plates has
been restored. (M. 9377.) } nat. size.
3. Crown view of the first and second right lower molars (or last milk-molar and
first molar) of Hlephas antiquus Falconer. (M. 9384.) 3 nat. size.
(The numbers of the specimens are those in the British Museum register.)
3. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904—-1905.—
Report on the Polyzoa. By Cuartes I. RoussELet,
F.R.M.S.
TReceived February 2, 1907. ]
(Plates XIV. & XV.*)
The freshwater Polyzoa collected in Lake Tanganyika by
Dr. W. A. Cunnington are attached to stones and shells which
were partly obtained in shallow water and partly dredged from
20 to 40 fathoms.
Altogether the collection contains five species, three of which
belong ‘to the Phylactolemata and two to the Gymnolemata.
Amongst the latter is Moore’s Arachnoidia ray-lankestert (107),
which was found in some abundance on shells of Paramelania
dredged from deep water.
Two species of the Phylactolemata are of the Plumatella type,
with horseshoe-shaped lophophore. One of these appears to be a
new species, very closely adherent to stones, with half-formed
ea-shaped tubes, which I have named Plumatella tanganyike.
To the second species I have given no name, as the few frag-
ments of tubes and the total absence of statoblasts offer no
characters that would distinguish it from Plumatella repens.
The third Phylactolematous specimen is an interesting new
species of the genus Fredericella, which I have named Fredericella
cunningtont in honour of its discoverer, who dredged it from
25 fathoms near Mshale.
The second of the Gymnolematous species is of special interest,
as being always found associated with, and imbedded in, a fresh-
* Wor explanation of the Plates, see p. 257.
+ The numbers refer to the Bibliography, pp. 256, 257.
IE ZS LO Tay
Fig. FRDixon-Nuttall. del. Highley, del et lith.
TANGANYIKA POLYZOA.
Nae
lee youll SOV Nell ONE
Highley, del et lith.
Fig 10. FER Dixon-Nuttali.del.
TANGANYIKA POLYZOA.
1907. ] OF THE THIRD TANGANYIKA EXPEDITION. 251
water sponge, Spongila tanganyike Evans, only the small heads
projecting beyond the surface of the sponge. Its affinities appear
to approach nearest to Victorella, and I have therefore named it
Victorella symbiotica.
The following is a list of the five species here described :—
Plumatella tanganyike, sp. n.
. repens, var, ?
Fredericella cunningtoni, sp. n.
Arachnoidia ray-lankesteri Moore.
Victoreila symbiotica, sp. n.
The number of known species of Polyzoa inhabiting African
fresh waters is thus brought up to eight, truly a remarkably
small number for this vast continent.
The other African species so far recorded are the following :—
Fredericella sultana, found by Dr. Stuhlmann in Alexandria,
Kgypt (7), and in the Equefa River, Natal, by the
Hon. Thos. Kirkman *,
Plumatella repens, var., from Ugogo, Victoria Nyanza, Albert
Edward Nyanza,and Albert N yanza : the statoblasts only
found by Dr. Stuhlmann (7) and also by Dr. Meissner
on shells of Wtheria (in the Berlin Museum) from the
Upper Nile, White Nile, the Niger, and Senegal (8, 9).
Lophopodella (Pectinatella) carteri Hyatt, from Ugogo:
statoblasts found by Dr. Stuhlmann (7).
Lophopodella thomasi Rouss., from Hunyani River, Rho-
desia (12).
The indifferent preservation of the Tanganyika specimens has
unfortunately hampered and prevented a complete study of all
the species. They were narcotised with cocaine and then pre-
served in alcohol, which is not a sufficiently good fixative for
these animals. Freshwater Polyzoa must not be allowed to die
in the anesthetic, which quickly macerates their delicate bodies.
After treatment with cocaine in perfectly clean water they should
be killed and fixed whilst still living, either with very weak
osmic acid (;'5 per cent.) for ten minutes to half an hour, according
to the age and actual strength of the solution, then washed, and
also preserved, in 3 per cent. commercial formalin (97 c.c. water,
3 c.c. formalin); or else fixed with a 10 per cent. solution of
formalin (90 ¢.c. water, 10 c.c. formalin) for 24 hours, then
preserved in 3 per cent. formalin to which 3 per cent. glycerine
may be added to render the animals more transparent. Polyzoa
fixed with osmic acid are stained brown, or even black, if allowed
to stay too long in the fixative; those fixed with formalin remain
white and transparent.
The little bottles in which the preserved specimens are stored
should have no air-space; an air-bubble plays havoc with the
delicate tentacles of extended polypides.
SUB Oo bo
* Mentioned in his paper on the Rotifera of Natal as supporting tubes of Limnias
ceratophylli: Journ. Roy. Micr. Soc. 1901, p. 232.
252 MR. C. F. ROUSSELET ON THE POLYZOA [ Mar. 5,
With regard to the question of the origin of the freshwater
Polyzoa, I am inclined to agree with Dr. Wesenberg-Lund (18),
who has expressed the view that all the different groups have
wandered from the sea, the Phylactolemata at an early period, so
that their marine ancestors are not now known, and may have
become extinct in the sea long ages ago, whilst the freshwater
Gymnolemata, i.e. Paludicella, Victorella, Pottsiella, Arachnowdia,
have immigrated at a much later period, so that they still show
some aftinity with several marine genera.
Tf this conception be correct, it follows that there can be no
relationship between the living Phylactolemata and Gymnolemata,
and still less can there be intermediate forms connecting the one
with the other.
In June of last year I succeeded, with the kind assistance of
Dr. Bousfield, in again finding Victorella pavida in the Surrey
Canal, London, where it had been obtained some twenty years
ago, but not recorded since. A study of this species and of the
other known freshwater Gymnolemata has impressed upon me
that, in addition to the absence of statoblasts, they are all pos-
sessed of a common character of considerable importance, which
they share with a group of marine Ctenostomata, and which may
well denote a certain degree of affinity. In all these forms there
is a stolon which expands into a cell or zocecium, at the upper end
of which an orifice is formed which may become enlarged into an
elongated, more or less cylindrical tube for the protrusion of the
polypide. Behind the orifice the stolon, after forming a septum,
continues to grow out to form another cell a little further on ;
then on each side of the cell normally one additional stolon arises,
also separated by a septum, to form new branches, which repeat
the same process of cell-formation until the growing point of
the stolon is either broken off or becomes atrophied. In this way
a zoarium is produced, forming an irregular network of branches,
approximately at right angles to each other. This cruciform
mode of growth can readily be observed in all the species of the
following genera, which might be grouped together under the
name of ‘ Cruciform Stolonifera ” :—
Paludicella.
Victorella.
Pottsiella.
Arachnoidia.
Freshwater.
J
Arachnidium. | nee
oe 3 + Marine *.
Cylindreciun, |
PLUMATELLA TANGANYIK#, sp.n. (Plate XIV. figs. 1—4.)
Specijic Characters—Zoarium consisting of clear light brown
chitinous tubes, branching, curving, and interlacing, closely
* The marine species Barentsia misakiensis from Japan, described by Dr. A. Oka
in 1895, shows the same fundamental structure.
1907. | OF THE THIRD TANGANYIKA EXPEDITION. 2S
adherent, encrusting stones, in the substance of which they are
partly embedded ; also encrusting shells of molluscs; tubes some-
times flat-sided. Tentacles about 20. Sessile statoblasts oval ;
floating statoblasts not observed.
The appearance of this new Plwmatella is very different from
the European species and varieties of the genus, as will be seen
by figs. 1 & 2, Pl. XIV.
The tubes form a thin, closely adherent, interlacing, encrusting
layer, and are partly embedded in the stone on which they grow ;
or, possibly, stony crystalline material has been deposited in
between and over the tubes. The tubes encrusting stones are
fairly tubular, but those encrusting molluscan shells are flat-sided
or e&-shaped in section, and the adhering side has only a very
thin layer of chitinous material, so that the supporting shell is
used by the animal to form part of its protecting tube. The
tubes are of a light brown colour, clear and nearly transparent.
The raised ends of the tubes are always tubular and white.
Septa are present in the tubes at the poimts of branching.
A full and well-extended head was not found; but there is
every appearance that the lophophore is horseshoe-shaped, with
about 20 tentacles.
Sessile statoblasts were found in the tubes (Pl. XIV. figs. 3 & 4);
they are oval in shape, but varying a good deal in their pro-
portions of width to length, smooth, and surrounded by a thin,
narrow, flat annulus without air-cells; their greatest size is:
length “450 ue (iy in.), width 343 Ge in.). Ordinary floating
statoblasts with swimming annulus were not observed.
Diameter of tubes 321 y (5 im.) on average.
Habitat. Encrusting stones just below water-level at Kalambo
and Kassangiu; and also encrusting shells in shallow water at
Chamkaluki.
PLUMATELLA REPENS, var. ¢
This specimen was found attached to shells of living 7iphobia,
dredged from about 40 fathoms at Maswa.
The tubes are brown in colour, semi-opaque, and show a
deposit of fine material in more or less regular longitudinal or
transverse lines; they are partly adherent, branching irregularly,
tubular, and altogether have much resemblance to those of
Plumatella repens of our lakes and canals, from which it is hardly
possible to differentiate them.
Fully extended specimens were not present, but the lophophore
appears to have about 22 to 24 tentacles.
The diameter of the tubes is 364 jy (,5 in.) on average.
Some sessile statoblasts found in the tubes are oval, smooth,
with thin flat annulus without air-cells, 407 (;4 in.) in length
and 278 (_; in.) im width. Floating statoblasts with cellular
annulus were ‘not observed.
The tubes are larger, and the sessile statoblasts smaller and
more elongated, than in the preceding species.
Proc. Zcou. Soc.—1907, No. X VITI. 18
254 MR. C. F, ROUSSELET ON THE POLYZOA [ Mar. 5,
FREDERICELLA CUNNINGTONI, Sp. n. (Plate XV. figs. 9 & 10.)
Specific Characters.—Zoarium consisting of creeping, closely
adherent, branching and interlacing €&-shaped tubes, built of a
very thin, transparent, chitinous, internal membrane, covered
externally with coarse grains of sand; lophophore circular, with
16 tentacles.
The appearance of the tubes of this new species is such that they
were at first passed over as apparently the tubes of some aquatic
worm or insect larva, until a circular lophophore was observed
protruding from one of the tubes, when a closer examination,
and the removal of the adhering tubes, revealed an abnormal
Polyzoon.
Though the indifferent preservation of the specimens did not
allow me to ascertain with certainty whether an epistome is
present or not, all the other characters are those of a phylacto-
lematous Polyzoon; and as the lophophore is circular, the genus
Fredericella naturally suggested itself.
The tube is low and €-shaped in section, very closely adherent
to the surface of shells and stones, branching and interlacing
freely. The structure of the tube is very abnormal; it consists
of a very thin, transparent, chitinous layer, to the external
surface of which grains of sand of various sizes and colour are
cemented, not unlike the tubes of some marine worms. The
grains of sand are mostly angular white quartz, and, to a less
extent, green olivine, interspersed with red and black grains, and
occasionally some plant and shell fragments. A few specimens
show tubes made of finer sand-grains. Only the raised ends of
tubes are tubular, otherwise the whole of the creeping tube is
@-shaped, the adhering surface being free from grains of sand.
The cells of the polypides are long, continuous, and with a few
incomplete septa here and there. The width of the tubes is
346 uw (4 m.) on an average.
The lophophore is circular and has 16 tentacles, but the rest
of the anatomy could not be made out, owing to the bad state of
preservation. No statoblasts were found in the tubes.
Hab. Encrusting stones and Weothawma shells dredged in
about 10 fathoms at Kibanga, and also shells dredged in about
25 fathoms near Mshale.
The genus /redericella has contained, so far, only one species,
i. e. #. sultana of world-wide distribution, if we consider the
three American forms /. regina, walkottti, and pulcherrima
as synonyms of /. sultana, in accordance with the opinion of all
recent authorities. /. duplessisi, found by Dr. F. A. Forel
(2, 3, 4) living unattached in the soft mud at the bottom of the
Lake of Geneva, also is but a /. swltana which has adapted itself
to its environment. Having examined specimens dredged by
Dr. Forel, consisting of short tubes with few branches, not
exceeding half an inch in length, I can find no difference between
them and the type species. J should like to mention here that I
find the tubes of /’, sultana rarely cylindrical, as has usually been
1907. | OF THE THIRD TANGANYIKA EXPEDITION. 255
stated; much more frequently they are distinctly angular, and
mostly triangular in shape.
ARACHNOIDIA RAY-LANKESTERI Moore. (Plate XIV. figs. 5 & 6.)
In his book ‘The Tanganyika Problem,’ 1903, Mr. J. E. 8.
Moore (10) mentions on p. 295 a gymnolematous Polyzoon found
by him on shells of Paramelania dredged from about 20 fathoms
off the shore of the lake, to which he gave the above name,
considering it allied to the marine genus Arachnidiwm.
This interesting species has been found again by Dr. Cunnington,
also on shells dredged from 20 to 25 fathoms near Mshale. The
colonies form an encrusting network, consisting of ovoid or
irregular, membranous, brownish cells, with toothed edges, closely
adherent to the shell, and connected together by narrow, tubular,
creeping stolons. The cells or zocecia are very flat and shallow,
and have at their upper end a tall erect tube from which the
polypides protrude. The lophophore is circular, without calyx,
and has 16 narrow tentacles. A septum can be seen at the origin
of each stolon.
In retraction the polypide retreats down the erect tube into
the cell, invaginating in the process the anterior thin-walled part
of the tube, which is the tentacle-sheath or kamptoderm. All
that can be perceived within the shallow cell is a band of retractor
muscles, and a sac filled with granules, possibly the stomach.
This species belongs to the group of Gymnolemata which I
have called “Cruciform Stolonifera,” and certainly appears closely
allied to the marine genus Arachnidium.
Its mode of growth appears to be as follows:—A narrow,
filiform, closely adherent stolon, following all the irregularities
of the surface of the shell, expands to form an ovoid cell, the
edges of which are more or less toothed, the better to fit the
depressions and ridges of the shell-surface. At the upper end
of the cell an orifice is formed which elongates into an erect tube,
whilst behind the orifice the stolon, after forming a septum,
continues to grow to form another cell at some distance further
on. At an early stage of the formation of the cells, stolons can
be seen growing out laterally, one on each side, rarely more, to
form side branches, and thus a very irregular cruciform network
is formed. The cells are always well separated from each other
by a stolon, and when found agglomerated together, as in
Mr. Moore’s rough sketch, it is the result of two or more colonies
having grown over and in between each other.
Of the polypide the indifferent preservation does not allow me
to say more than that the lophophore has 16 narrow tentacles.
The size of the largest cell seen is 792 4 (34 in.); width of
stolon 69 (s+ > im.); length of the tube when the animal is
protruded 970 (4; im.).
VICTORELLA SYMBIOTICA, sp.n. (Plate XV. figs. 7 & 8.)
Specific Characters—Zoarium a narrow, tubular stolon with
18*
256 MR. C. F. ROUSSELEL ON THE POLYZOA [ Mar. 5,
slightly expanded portions at imtervals, from which a long erect
tube rises; side branches arise at the base of the zoecia.
Polypides with cireular lophophore of 8 tentacles. The whole
goarium and zocecia embedded in Spongilla tanganyike, on the
surface of which the polypides protrude.
The freshwater sponges brought back by Dr. Cunnington
from Lake Tanganyika include specimens of Spongilla tanganyike
Eyans, in which was found embedded this gymnoleematous polyzoon
which cannot be identified with any known species. _ Its affinities
seem to lie between the genera Cylindracium, Pottsiella, and
Victorella; but its habitat in freshwater and the number of
tentacles possessed by the polypides, have decided me to place it
in the last-named genus.
Potts’s Paludicella erecta (11), afterwards renamed Potisiella
erecta by Prof. Kraepelin (6), is the only other freshwater species
which is known to penetrate encrusting sponges, but its circular
lophophore has 20 tentacles.
The meandering cylindrical stolons, glassy white when cleared
from adhering sponge-fragments, form an entangled mass difficult
to follow, and are entirely embedded in the substance of the
sponge. Adjoining and crossing stolons and tubes adhere, but
can be separated by tearing apart. No septa have been observed
in the stolons.
The upright tubes, together with the slight expansions of the
stolon, form the cell or zocecia, and lateral branches, one on each
side, were observed springing from the expanded stolon, but these
side branches are not always present.
The tubes are long, erect, single, slightly widened at the base,
of glassy transparency, except the small portion above the surface,
which is rendered more or less opaque by fine granules, and
emerge on the surface of the sponge where the polypides expand
their small circular lophophore of 8 tentacles. The length of the
tubes in this species is no doubt due to the necessity of reaching
to the surface of the sponge. In Victorella pavida of Kent (1, 5)
the cell is formed by a very distinct expansion of the stolon and
the tube arising therefrom is of moderate length. Length of
erect tubes up to 1:7 mm. (;/5 in.).
It has been a matter of considerable difficulty to dig the
complete stolon with the tubes out of the substance of the sponge
without breaking up these delicate structures, and the operation
has been only partially successful.
Hab. Growing within and through specimens of Spongilla
tanganyike, which were found encrusting rocks in shallow water
at Ghamkaluki, and also on shells dredged from about 20 fathoms
near Mshale.
Bibliography.
1. Bousrretp, E. C.—The Victorella pavida of Saville Kent.
Ann. Mag. Nat. Hist. ser. 5, vol. xvi. 1885, pp. 401-407,
1, plate.
1907.| OF THE THIRD TANGANYIKA EXPEDITION. 257
2.
10.
ihe
12.
13.
Fig.
Fig.
Du-Puessis-Gouret, Dr. G.—Essai sur la faune profonde des
lacs de la Suisse. Mém. Soc. Helvétique d. Sc. Nat.,
vol. xxix. Bryozoaires. Genéve, 1885.
Foren, Dr. F. A~-Faune profonde des lacs Suisses. Meém.
Soc. Helvétique d. Se. Nat., vol. xxix. Bryozoaires.
Geneve, 1885.
——. Le Léman. Lausanne, 1904, vol. iil. pp. 115-114, 300.
Kent, W. 8.—On a new Polyzoon, Victorella pavida, from the
Victoria Docks. Quart. Journ. Micr. Se. new ser. vol. x.
1870, pp. 34-39, 1 plate.
Krarpretin, Prof. Dr. Karu.—Die Deutschen Siisswasser-
Bryozoen. Hamburg, 1887.
. Metssner, M.—Moosthiere. Deutsch-Ost-Afrika, Band iv.
1895.
Beitrag zur Kenntnis der geogr. Verbreitung der
Bryozoengattung Plumatella in Afrika. Zool. Anz. no. 430,
1893.
Weiterer Beitrag zur Kenntnis der geogr. Verbreit-
ung der Siisswassergattung Plumatella. Zool. Anz. no. 531,
1897.
Moore, J. EK. $.—The Tanganyika Problem. London, 1903.
Arachnoidia, p. 295.
Ports, Epwarp.—On Paludicella erecta. Proc. Ac. Nat. Se.
Philadelphia, vol. 1. 1884, pp. 215-14. Reproduced also in
Ann. Mag. Nat. Hist. ser. 5, vol. xiv. 1884, pp. 437-439.
Rovussevet, Cuas. F.—On a new Freshwater Polyzoon from
Rhodesia (Lophopodella thomas). Journ. Quekett Mier.
Club, ser. 2, vol. ix., April 1904, pp. 45-56.
WESENBERG-LuND, Dr. C.—Biologiske Studier over Fersk-
vandsbryozoer. Kj@benhavn, 1896, 4 plates. Résumé en
francais.
EXPLANATION OF THE PLATES.
PrarEe XIV.
. Plumatella tanganyike (p. 252). Portion of interlacing tubes. X 15.
Plumatella tanganyike. Flat under surface of tubes detached from shell,
showing septa. X Jd.
. Plumatella tanganyike. Sessile statoblast, upper surface. X 50.
. Plumatella tanganyire. The same, under surtace. 50.
. Arachnoidia ray-lankesteri (p.255). Colony growing on shell. X 20.
. Arachnoidia ray-lankesteri. Single cell and polypide. X 60.
Qe poe
PratTe XV.
7. Victorella symbiotica (p. 255). Piece of sponge with polypides protruding
on surface. X 20.
8. Victorella symbiotica. Cells and stolon separated from sponge. 20.
9. Fredericella cunningtoni (p. 254). Interlacing tubes adhering to shell
surface. X 15.
10. Fredericella cunningtoni. Portion of same with protruding polypide.
x 56.
258 DR. W. A. CUNNINGTON ON THE BRACHYUROUS [ Mar. 5
4. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905.—
Report on the Brachyurous Crustacea. By Wui.tiam
A. Counnineton, B.A, Phe D., F.Z:S:
{Received March 5, 1907. |
(Plates XVI. & XVII.* and Text-figure 84.)
1. Lntroduction.
Since the year 1896, when Mr. J. EH. 8S. Moore paid his first
visit to Lake Tanganyika, our knowledge of the fauna of that
lake, and of the interesting problems which are connected with it 7,
has considerably advanced. The single representative of the
Brachyura then collected was described somewhat later t, but
already, in 1887, a description had been given by Milne-Edwards §
of another form brought from the lake by Captain Joubert. The
description of this last-mentioned type was unfortunately very
incomplete, while at that time no male specimen had been
obtained. During Mr. Moore’s second expedition, in 1899-1900,
he succeeded in collecting further examples of both the species
already known, and furnished male specimens of the form which
Milne-Edwards had described from the female alone. Finally, a
Third Tanganyika Expedition was dispatched in 1904, with the
conduct of which I had the honour of being entrusted, and which
has added yet again to our knowledge of the Brachyurous
Crustacea of the lake.
The very earliest facts which were learned respecting the fauna
of Tanganyika seemed to show that the animals it contained
were not merely of an unusual type, but were different from those
living in the other big African lakes. It is an interesting fact
that, with the lapse of time and a very great increase in our
knowledge of the lakes of Central Africa, this dissimilarity of
Tanganyika has been not only confirmed, but rendered more and
more striking. Among other animal groups, that of the Brachyura
affords an excellent example of this remarkable state of things ;
so that a double purpose will be served by the description in this
paper of Crabs from Nyasa, which lake has the normal charac-
teristics of the African fresh-waters.
The collection of Crabs made during this Expedition contains
representatives of five species, of which two alone have been
previously described. By the kindness of Dr. W. T. Calman, I
have been permitted to examine and compare with the collection
a large number of specimens of nearly allied forms, belonging to
For explanation of the Plates, see p. 276.
Moore, ‘ The Tanganyika Problem ’ (London, 1903).
Cunnington, Proc. Zool. Soc. 1899, p. 697.
Ann. Sci. Nat. 7° ser., Zool. t. iv. (1887) p. 146.
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1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 259
the British Museum *. Among others are some which are stated
to have come from Nyasa and Tanganyika; and since no notice of
these specimens appears to have been published, it seems desirable
to consider them, also, in the present paper. The following is
thus a complete list of the species at present known to occur in
the lakes in question :—
Nyasa.
Potamon (Potamonautes) inflatus (H. Milne-Edwards).
¥5 (Potamonautes) orbitospinus, sp. i.
i" (Potamonautes) sp. ?
TANGANYIKA.
Potamon (Potamonautes) platynotus, sp. n.
“ (Potamonautes) sp. ¢
Platythelphusa armata A. Milne-Edwards.
maculata (Cunnington).
conculcata, sp. 0.
99
2?
2. Systematic Notes and Description of New Species.
Family PoTAMONID# (= THELPHUSID 2).
Subfamily Poramontn a.
Lake Nyasa.
Poramon (Poramonautes) tNFLATUS (H. Milne-Edwards).
Potamon (Potamonautes) inflatus Rathbun, Nouv. Arch. Mus.
Hist. Nat. Paris, 4° sér., vil. (1905) p: 174 (ubi synon.).
A single fair-sized male specimen in the collection of the
British Museum (Reg. No. 96.7.19.1) appears to belong to this
species. The only particulars given are :—“ Kavisonda, Nyasa.
Presented by Dr. Ansorge.”
PoraMon (POTAMONAUTES) ORBITOSPINUS, sp. n. (Plate XVI.
fig. 1.)
Description.—Carapace considerably flattened, antero-lateral
margins arcuated and denticulate, extending laterally less than an
orbit’s breadth beyond the external orbital angle. No definite
epibranchial tooth, but the end of the post-frontal crest forming
a distinct corner at that point. Regions and sutures on carapace
moderately marked, lateral regions exhibiting a series of small
slightly oblique granular ridges. Front rather deflexed, less than
one-third the width of the carapace, and with its anterior margin
* By the courtesy of the authorities of the Berlin Museum, co-types of some of
the species described by Hilgendort from German Hast Africa were lent to the
British Museum for the purpose of comparison with those discussed here. I am
especially indebted to Dr. P. Pappenheim for detailed information regarding these
specimens.
260 DR. W. A. CUNNINGTON ON THE BRACHYUROUS — [ Mar. 9,
sinuated. _Post-frontal crest prominent, almost straight, and
extending to margins, with branches of mesogastric groove
angulated. Orbits large, and eyes large, with stout peduncles.
External orbital angle produced into prominent spine. Ischium
of external maxillipeds showing longitudinal furrow, somewhat
nearer to the inner edge. Chelipeds in both sexes subequal ;
merus trigonous, with a series of small’ spines and distally a pro-
minent spine on the anterior margin; carpus with two spines on
inner margin, the posterior being the smaller. Fingers pointed
and slightly hooked; teeth fairly uniform, of moderate size.
Ambulatory legs long and slender, also somewhat compressed.
Colour in life dull dark green, shading into dark purple; the most
prominent parts of the front, the external orbital spines, antero-
lateral margins, and portions of post-frontal crest outlined in
white; legs dark purple.
Dimensions as follows :—
Adult male (largest specimen) :
mm.
Length of carapace ............ 384
Breadth of carapace ............ 56°9
Fronto-orbital width ......... 36°7
NAAUGUG, GL THROWING sun odeos somone oa 13°8
Adult female :
eneth of carapace 22.5)... 36°9
Breadth of carapace ............ 53°9
Fronto-orbital width ......... 34°5
AY AUCHEOL GE THEOL ad cddscddcocsene st 14-2
Remarks.—As there has recently been published a very com-
prehensive revision of the Potamonide by Miss Rathbun %, it
becomes desirable to take this asa basis, and correlate with it any
newly described forms. Accordingly that portion of the key
given by Miss Rathbun’? which is affected is restated below, but
slightly modified to include this new species, while distinguishing
it from johnston, the species close to which it finds a place.
h'. Carapace broader; less than three-quarters as long as broad.
Jj. Antero-lateral margins denticulated; external orbital angle
produced into prominent spine saa pecans aeeodnG orbitospinus.
j'. Antero-lateral margins granulated; external orbital angle
forms low blunt process ..............0...000ceetereeeccceeeeeess es Johnstoni.
Even a casual study of the species comprised in the genus Potamon
renders it abundantly evident that in many cases they cannot be
distinguished by very striking differences of character. It is
therefore the more important to make clear the actual points of
disagreement which have been thought in this case to justify the
formation of a new species. PP. orbitospinus approaches most
* Nouv. Arch. Mus. Hist. Nat. Paris, 4° sér., vi.—viii., 1904-1906.
+ Op. cit. vii. p. 160.
1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 261
nearly to the forms P. hilgendorfi (Pfeffer) and P. johnstoni
(Miers). In order to render a comparison more easy, the
diverging characters of these three species are given in tabular
form, while it may here be stated that P. orbitospinus is a species
certainly more easily recognised than some of those which are
allied to it.
hilgendorfi (Pfetf.)
(not hilgendorfi Hilg.). TALMOS POPS SDs Le
johnstoni (Miers).
Carapace somewhat flat- | Carapace considerably
tened. | flattened.
I
Antero-lateral margins | Antero-lateral margins
granulated. denticulated.
External orbital angle | External orbital angle pro-
} . .
rather acute. | duced into prominent
spine.
Fronto-orbital width °71 | Fronto-orbital width *64
that of carapace. | that of carapace.
Ischium of external maxil- | Ischium of external maxil-
lipeds without longi-| lipeds with longitudinal
tudinal furrow. | furrow.
Posterior carpal spine | Posterior carpal spine (of
(of cheliped) little | cheliped) sharp and
developed. | well developed.
| Ambulatory legs long and
Carapace somewhat flat-
tened.
Antero-lateral
eranulated.
margins
External orbital angle
forms low blunt process.
Fronto-orbital width °59
that of carapace.
Ischium of external naxil-
lipeds with longitudinal
furrow.
Posterior carpal spine (of
cheliped) little deve-
loped.
Ambulatory legs short
and stout.
| slender.
Occurrence.—N kata Bay, 23/6/04. Several specimens of medium
size. The Crabs were captured and brought me by the natives,
so that I have no direct evidence of their mode of life. They are
said to live in the water and not on land, though they sometimes
come on to the beach.
Three small and immature specimens in the collection of the
British Museum (Reg. No. 91.12.19.1-3). Only the following
particulars are given :—‘‘ Lake Nyasa, Coll. Miss M. Woodward,
pres. by Miss Sophia McLaughlin.”
One rather small male, no precise locality given, British
Museum (Reg. No. 93.1.14.1). ‘“‘Lake Nyasa. Received from
Mr. Joseph A. Williams, Universities Mission, Likoma, Lake
Nyasa.”
One large female specimen, British Museum
97.4,29.1).
Bay to Ruarwe, June 1896.”
H. H. Johnston.”
A single large male specimen, collected by Mr. J. E. 8. Moore
during one of his expeditions to Tanganyika, but without further
particulars.
(Reg. No.
“West coast of Lake Nyasa from Ukala (? Nkata)
“A. Whyte collector—pres. by Sir
262 DR. W. A. CUNNINGTON ON THE BRACHYUROUS _[ Mar. 9,
Poramon (POTAMONAUTES) sp. @
Reference has already been made to the fact that it is often
difficult to distinguish between closely similar species of this
genus. Many of the characters enumerated by the systematic
writers on this group, while in certain cases affording a ready
means of separating species, are in other cases of little or no
value. A careful consideration of many of the described species
and the examination of an extensive series of specimens lead to
the undoubted conclusion that some species have been established
without sufficient justification. While a species or even a genus
may be established on the strength of a single specimen, if its
characters are sufticiently unusual, it is necessary to be very
cautious in doing any thing of the kind within the limits of this
subgenus (Potamonanutes) in particular. Yet there are several
species of Potamonautes based upon single specimens, and even
upon specimens which were recognised as being immature. We
do not know a great deal about the modifications in form produced
by advancing age, but we do know, from the examination of any
extensive series, that they are considerable. The proportion
of length of carapace to its breadth, with the relative proportions
of the front, the orbits, and the fronto-orbital width—all of
them characters employed by the systematist—are undoubtedly
dependent upon the age and growth ‘of the individuals.
But more than this. A natural hesitation to lay stress on
every little difference in form is only emphasized by the result
of Schenkel’s* investigation of the species Potamon (Potamo-
nautes) celebensis de Man. He is able to distinguish some six
local varieties, in addition to the forma typica of de Man. These
differ from one another in colour, in the shape of legs and carapace,
in the extent of the sculpturing r, and especially as regards the
degree to which the carapace is vaulted and the br omental regions
dilated. But, as Schenkel points out, these features are precisely
those which would be affected by a difference of surroundings.
He finds the Crabs with the flattest carapace come from mountain-
streams, where of course the water is pure and well oxygenated.
The converse holds equally good, the gill-chambers in other
varieties being inflated in proportion to the sluggish or muddy
nature of the river inhabited, and it is obvious that the sculpturing
is In a great Measure an expression of the degree of inflation of
the carapace. Again, it is pointed out that the amount of the
food-supply must have a powerful influence on growth: thus with
two Crabs of similar size, that from water poor in nourishment
will be far older than the other, and so not strictly comparable
with it.
As Schenkel considered it desirable to retain a single species,
but to constitute a number of varieties in this case, this discussion
of the facts is very suggestive in a general sense. Under such
circumstances, some observers would not even separate into
* Verh. naturf. Ges. Basel, Bd. xiii. 1902, p. 528.
1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 263
varieties, and, on the other hand, there can be no doubt that many
so-called species of other writers are not more distinct than these
varieties of Schenkel. If it is possible to trace a gradual
transition from one variety to another, as appears to be the case
with the form celebensis, it becomes evident that certain established
species are of similarly little value, and that species of this genus
ought never to be constituted on such slender grounds.
This matter is dealt with at such length, because the conviction
has forced itself upon me that the number of African species, at
all events, has been multiplied to an excessive degree. Where
two or more well-marked species exist side by side, there can be
no hesitation in the matter; but our knowledge of the Crabs of
the African continent is still very scanty, and over a vast area
restricted to a few isolated records. Thus it is impossible to
determine yet whether this or that species may not prove to be only
a slight local modification of some widely distributed form. This
being the case, I have not ventured to name the two specimens
from Nyasa, which are placed under the head-line above.
Although it was during my study of the two species described
as new in this paper that I became impressed with the unsatis-
factory character of some species of Potamonautes, it was not
until [ examined these two Nyasa specimens and a third from
Tanganyika that [found a case where the most careful comparison.
only resulted in confusion with several closely allied species. The
two specimens from Nyasa—a rather small male and a small
female—are in the collection of the British Museum (Reg. No.
97.4.29.2-3), and the following note is given as to locality :—
“ Kondowe to Karonga, N. Nyasa, June 1896. A. Whyte
collector—pres. by Sir H. H. Johnston.”
Using asa basis the key in Miss Rathbun’s recent revision,
we find they belong to group c, where the lateral margins extend
less than an orbit’s breadth beyond the external orbital angle.
Further, they would seem to belong to division e’, where the
post-frontal crest is little advanced in the middie, and to sub-
division f', with fingers of the cheliped but slightly gaping.
Thus they would come nearest to the species hilgendorfi ( Pfeff.) and
johnstoni (Miers). The specimens differ from hilgendorfi (Pfett.)
principally in possessing a furrow on the ischium. From johnstont
they differ in being distinctly longer in proportion to the breadth,
in the sculpturing being less marked, and in the absence of the
fairly stout spinules on the anterior margin of the carpus of the
cheliped—a single distal spine only being present.
They agree to some extent also with suprasulcatus pseudo-
perlatus, which, if it exist at all as a true variety, is In a Measure
intermediate between hilgendorfi (Pfeft.) and johnstont. The
length in proportion to the breadth is, however, more extreme in
the case of suprasulcatus pseudoperlatus. From another important
type in group c—perlatus itself—these individuals do not differ
greatly, the carapace being of much the same proportions as
regards length and breadth. The width of the front is, however,
264 DR. W. A. CUNNINGTON ON THE BRACHYUROUS [Mar.5,
considerably less than in perlatus, and the post-frontal crest does
not extend forward so far in the middle. The posterior carpal
spine of the cheliped is also sharper and more prominent. Both
the male and female specimens show only a single transverse
furrow on the sternum in front of the abdomen, while in the
specimens of allied species examined the males almost invariably
have two furrows even in young individuals.
From this description of the characters of these two Nyasa Crabs,
it will be evident that my former contention is not without
foundation. These individuals agree to a considerable extent with
several species, and where they disagree with one they agree with
another, while, to my mind, they have not a single character
which satisfactorily distinguishes them from those forms to which
they are undoubtedly allied. In the present state of our know-
ledge, it would be futile to attempt to determine which species
should be eliminated, and Miss Rathbun has certainly taken the
vight course in retaining most of the species hitherto described,
though it seems to have led her into some difficulties respecting
her key. Bearing in mind the facts which have been detailed at
some length above, I prefer, in a case like this, to leave the
specimens unnamed, and so at least avoid adding to the already
existing superfluity.
Lake Tanganyika.
PoraMon (POTAMONAUTES) PLATYNOTUS, sp. n. (Plate XVII.
figs. 1 & 3.)
Description.—Carapace much flattened, antero-lateral margins
aveuated and denticulate, extending laterally shghtly more than
an orbit’s breadth beyond the external orbital angle. No definite
epibranchial tooth, but the end of the post-frontal crest forming
a distinet corner at that point. Regions and sutures on carapace
ill-defined, postero-laterally a few inconspicuous granular ridges.
Front rather deflexed, less than one-third the width of the cara-
pace, and with its anterior margin sinuated. Post-frontal crest
prominent, slightly arcuated, but sinuated laterally, extending
to margins; branches of mesogastric groove straight. Orbits,
eyes, and peduncles small. External orbital angle existing as a
short blunt spine. Ischium of external maxillipeds not showing
longitudinal furrow, though this may be faintly mdicated in the
oldest specimens. Anterior portion of the sternum hairy in the
female, not hairy in the male. Chelipeds in both sexes subequal ;
merus trigonous, with a series of granules, and distally a stout
spine, on the anterior margin; carpus with a prominent spine on
the inner margin and a slight process just behind it. Fingers
distally hollowed out, spoon-shape and meeting closely in a sharp
cutting-edge (Plate XVII. fig. 3); dactylus usually slightly
longer than the pollex. Teeth fairly uniform, of moderate size,
but the fingers of the larger chela gape a little and have proximally
a few larger flat crushing-teeth. Ambulatory legs of moderate
length, little compressed. Colour in life a uniform dull greenish
brown ; the dactyli of the chelipeds in the males usually black.
1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 265
Dimensions as follows :—
Male (probably young specimen) : mm.
Length of carapace............-.. 20°3
Breadth of carapace ...........- 30°7
Fronto-orbital width ............ IS 7
ANVGWGKGli it MNROVMG “gececbeseeovscoco: 10-1
Adult female (largest specimen) :
Length of carapace ............ 33°7
Breadth of carapace ...........- 48-2
Fronto-orbital width............ 30-0
VivatGlelhh Ohi TORO, Coo gecAeecooeBanns 13°83
Remarks.—This species comes into Miss Rathbun’s key under
the heading c', where the carapace extends laterally more than
an orbit’s breadth beyond the external orbital angle. It finds its
place in the same subdivision (g') as the species ambiguus and
mrogoroensis, and that portion of the key, modified to include it,
is accordingly given below :
g'. Anterior branch of cervical suture absent.
h. Ischium of external maxillipeds without longitudinal furrow.
;, Anterior portion of female sternum hairy................... 0 | latynotus.
3 JOM
j’. Anterior portion of female sternum TOG |NEVVO cosees _... Immrogoroensis”™.
7’. Ischium of external maxillipeds with longitudinal ftrrow... ambiguus.
The principal points in which platynotus differs from ambiguus
on the one hand and suwprasulcatus on the other are also given in
tabular form. The most striking features of the new species are
suprasulcatus Hilg. | platynotus, sp. Nn. ambiguus Rathbun.
Carapace moderately vaulted. Carapace much flattened. Carapace distinctly vaulted)
Antero-lateral margins granu- Antero-lateral margins Antero-lateral margins
lated. _ denticulated. granulated.
Anterior branch of cervical Anterior branch of cer- Anterior branch of cervi-
suture present. vical suture absent. cal suture absent.
Fronto-orbital width °60 that Fronto-orbital width °62 | Fronto-orbital width °57
of carapace. | that of carapace. that of carapace.
Ischium of external mavxilli- Ischium of external | Tschium of external maxil-
peds without longitudinal maxillipeds without lipeds with longitudinal
furrow. | longitudinal furrow. furrow.
|
Fingers pointed and slightly Fingers distally hol- | Fingers pointed and
hooked. lowed out spoon-shape. shghtly hooked.
the flattened carapace, the denticulated antero-lateral margin, and
the character of the fingers.
* Tt is by no meaus obvious why Miss Rathbun has inserted the form mvrogoro-
ensis here, under the heading c’, when it is quite clear from Hilgendort’s description,
266 DR. W. A. CUNNINGION ON THE BRACHYUROUS | Mar. 5,
Occurrence.—Kasakalawe and Kituta Bay, both south end of
Tanganyika. Several specimens; some females probably full-
grown, but the largest male apparently not so. The Crabs were
taken under boulders about high-water level.
Poramon (PoTAMONAUTES) sp.
Under this heading, I placea single rather small male belonging
to the British Museum (Reg. No. 89.2.8.1), of which the only
particulars are: “ Lake Tanganyika, E. C. Hore (ex coll.).” This
specimen agrees with and differs from its near allies in very much
the same way as the unnamed forms from Nyasa, which are
dealt with above. Thus it finds its place near to the latter, with
which it agrees in showing only a single transverse furrow on the
sternum in front of the abdomen, although a male. It disagrees
with the Nyasa form, however, in being less inflated and in
showing the sculpturing more distinctly. Further, the antero-
lateral margins are more finely perlated, and the spime on the
merus of the chelipeds is longer and sharper. In this case also
the specimen differs little from several species, and appears to
have no satisfactory distinguishing characters of its own, so that
I again take the course least open to objection and leave it
unnamed.
Genus PLuaryTtHEeLpHusSA A. Milne-Edwards.
Platythelphusa A. Milne-Kdwards, Ann. Sci. Nat., Zool. 7° ser.,
t. iv. (1887) p. 146.
Platytelphusa Hilgendorf, Deutsch-Ost-Afrika, Bd. iv. Lief. ix.
(1898) p. 21.
Limnothelphusa Cunnington, Proc. Zool. Soc. 1899, p. 698.
Limnothelphusa and Platythelphusa Moore, ‘The Tanganyika
Problem’ (London, 1903), pp. 280, 286.
Platythelphusa and Limnothelphusa Rathbun, Nouv. Arch.
Mus. Hist. Nat. Paris, 4¢ sér. vii. (1905) pp. 268, 269.
There area number of reasons which have led me to offer here
a new description of this genus established by A. Milne-Edwards in
1887. AsI have stated elsewhere *, the account given of the type
species, P. armata, and particularly the figures of it, leave a good
deal to be desired. Miss Rathbun reproduces a photograph of
this same individual, and adds something to the description; but,
as a result of the Second and Third Tanganyika Expeditions, we
very incomplete though it is, that his specimen should come into the group e¢, in
which the carapace extends laterally less than an orbit’s breadth beyond the external
orbital angle. While it may perhaps be doubted whether the distinction given
above under j and j’ is of much weight, it will be evident that if the species
mrogoroensis were withdrawn from its false position, the new form platynotus
would then be distinguishable from ambiguus by the difference of the external
maxillipeds. It is not necessary to discuss here the position which mzrogoroensis
should really occupy, the simpler course is taken of merely incorporating the new
species in the existing key.
* Proc. Zool. Soc. 1899, p. 701.
1907. | CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 267
now possess in this country a fairly complete series of this species
of both sexes. We have the possibility for the first time of com-
paring males with the female on which the genus was founded,
and this might in itself justify a redescription. Buta comparison
between a large series of the form described as Limnothelphusa
maculata and the specimens of Platythelphusa armata (which I
was unable to make in 1899), has convinced me that the former
cannot be regarded as constituting a separate genus, but falls mto
place as a species of Platythelphusa. A third species of Platy-
thelphusa is among the acquisitions of the last expedition, so that
the description which follows has been materially modified in
view of our much more extensive knowledge.
Description.—Carapace almost quadrilateral ; antero-lateral
margins arcuated and armed with spines ; postero-lateral margins
but shightly arcuated., Front little deflexed, nearly straight. Post-
frontal crest distinct and perlated, but not extending to lateral
margins. Sub-orbital spine more or less distinct, in addition to
prominent mner sub-ocular tooth. A stout triangular process
descends from the external angle of the front, and may be pro-
duced into a small spine antero-distally. Antenne situated partly
behind and partly between this descending process of the front
and the inner sub-ocular tooth ; the distal segments thus escape
an appearance of displacement by the front. Merus of external
maxillipeds broader than long, the palp being attached to its
antero-internal angle; ischium without longitudinal furrow.
Ambulatory legs somewhat compressed, the fourth leg consider-
ably shorter than the others, and with its two terminal segments
broad and flattened.
Remarks.—With the inclusion of Limnothelphusa, this genus
has lost nothing of its original distinctiveness. In the generic
description of Limnothel, phust, stress was laid on the simple 1 nature
of the second antennal segment, which was undistorted by the
deflexed front. Although it was perhaps not very happily ex-
pressed, this character is just as typical of the other two species
we know from Tanganyika, and deserves emphasis accordingly.
In those genera and subgenera where the front is considerably
depressed, the antenne have the appearance, at least, of a lateral
displacement or distortion in consequence. All the species of
Platythelphusa show the front little deflexed, but have a trian-
gular process descending from the external angle, which process,
however, passes to the front and side of the antenna, without
modifying its shape or direction. In order to make this distinctive
feature quite clear, the frontal region of all three species of Platy-
thelphusa is figured, while corresponding figures are given of
certain well-known species of the subgenera of Potamon (Plate X VI.
figs. 2-7).
Owing to the modification of the generic characters of Platy-
thelphusi and the suppression of ‘the genus Limnothelphusa,
Miss Rathbun’s key to the subfamily Potamoninz * requires
* Op. cit. t. vi. p. 245.
268 DR. W. A. CUNNINGTON ON THE BRACHYUROUS — [Mar.95,
alteration. In the restatement which follows, the subgenera are
omitted, as they are not affected, and care has been taken to
retain the original form as far as possible.
Subfamily PoraMonin”.
e. Ocular peduncles large, not tapering towards distal extremity.
f. A stout triangular process descends from the external
angle of the front ............. PLATYTHELPHUSA.
S'. No process desuanding Bunn the axiom anala a the
front.
g: Front not armed with spines or spinules ................... Poramon.
g'. Front armed with spinules ............ HyYDROTHELPHUSA.
. Ocular peduncles small, tapering tow andes ictal: amenity, TERIMETOPUS.
Since there are now three species included in the genus Platy-
thelphusa, 1t may be well to furnish a key to them, although they
are much more distinct and well defined than is the case with
many species of Potamonautes.
Key to the Species of PLAYYTHELPHUSA.
a. Carapace extending laterally more than an orbit’s breadth beyond
the external orbital angle; front less than one-third width of
carapace .........- . armata.
a’. Carapace exionitine TleGeonetlly: Tees ann an + ors s ineadihe he dnd
the external orbital angle; front more than one-third width of
carapace.
b. Carapace moderately convex; carpus of cheliped without spines
on outer margin; ambulatory legs of moderate length, anterior
margin of the merus without spines ........ .. maculata.
b'. Carapace extremely tlattened ; carpus of ghelipadl Theanine spines
on outer margin; ambulatory legs long, anterior margin
of the merus produced distally into two spines..................... conculeata.
PLATYTHELPHUSA ARMATA A. Milne-Edwards. (Text-figure 84.)
Platythelphusa armata A. Milne-Kdwards, Ann. Sci. Nat.,
Zool. 7° sérv., iv. (1887) p. 147.
Platytelphusa armata Hilgendorf, Deutsch-Ost-Afrika, Bd. iv.
Lief. ix. (1898) p. 22.
Platythelphusa armata Moore, ‘The Tanganyika Problem’.
(London, 1903), p. 286.
Platythelphusa armata Rathbun, Nouv. Arch. Mus, Hist. Nat.
Paris, 4° sér., vil. (1905) p. 269.
Description.—Carapace moderately convex, extending laterally
more than an orbit’s breadth beyond the external orbital angle.
The antero-lateral margins differ little in length from the postero-
lateral, with which they are almost directly continuous. Number
of spies on antero- lateral mar ein extremely variable, usually
four or five, in addition to the spine at the external orbital angle.
Regions and sutures fairly well marked. The postero- lateral
regions exhibit a series of small slightly oblique granular ridges.
1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 26%
Front less than one-third the width of the carapace, its margin
perlated, and with a stout spine at each extremity. Sub-orbital
spines short and stout; a small spine on the descending process
of the front. Orbits small, -16 width of carapace ; orbital margins
perlated. Eyes and peduncles small. Chelipeds in both sexes
unequal; merus trigonous, with distally a prominent spine on
the anterior margin, and a small spine or tubercle on the ventral
margin ; carpus with two spines on inner margin and one above
the point of articulation with the hand. Hands and fingers
somewhat compressed, the dorsal border being keeled and granu-
lated ; the ventral border in the larger chela characteristically
arcuated at the junction of the pollex. Fingers pointed and
slightly hooked ; those of the larger chela gape, and have a pair
of large flat crushing-teeth proximally, with a series of rather
smaller ones distally. In the smaller chela the teeth are fairly
uniform and of moderate size. Ambulatory legs of moderate
length. Colour, in life, yellowish brown, with irregular blotches
of darker brown on the carapace.
Text-fig. 84.
Platythelphusa armata, large male. X 3.
Milne-Edwards and Miss Rathbun give the detailed dimensions
of an adult female (type specimen), but as we now possess for
the first time particulars of males, the measurements of an
adult male—the largest known specimen—are included here.
In this individual (text-fig. 84) the size of the larger chela is very
striking, the length of the hand and pollex being greater than the
whole breadth of the carapace.
Proc. Zoot. Soc.—1907, No. XIX. 19
270 DR. W. A. CUNNINGTON ON THE BRACHYUROUS [| Mar. 5,
Adult male (largest specimen), Moore’s collection :
mm
Wenethvoncanapace werner arrears eee re rae eC ere 53°2
TereeAVGlilN Olt (CARGAORIE®, 9 Ghoasosv onvaddacboceubanvencoacades 66°3
JASN H-COre OMA VAMGHEIO papaccoopeseasdosodonacconnponcoouse 40-1
Waadth Of fromb Vili cores. een esc ae atone eee 18-7
Larger chela: greatest length of hand and pollex. 71:3
Pt ” greatest height of hand ............ 35°8
Remarks.—While the chelipeds in both sexes are unequal, it
appears that either the right or left may be the larger, quite
indiscriminately. The large crushing-teeth are often considerably
worn in old specimens, and the spines of carapace and chelipeds
become blunt and rounded. ‘This is, in fact, the case with the
male specimen which is figured, where it is clear that certain of
the spines have lost their original sharpness, but it should be
understood that the spines in question are normally very sharp
and strong. ;
A curious feature of the specimens of this Crab is the number
of circular blotches which appear on the surface of the carapace
and appendages. The marks are approximately round, and seem
due to an eating-away of the calcareous matter of these spots,
which occur in greater numbers on larger and older specimens.
In some cases the fingers of the chele have suffered severely,
being partially eaten through by this process of erosion. It
seems most probable that these blotches are due to the action
of boring Alge. A portion of test including such a spot was
decalcified, cleared, and mounted in balsam, but showed little
structure even then. From a central point, a number of fine
processes, or perhaps tubules, could be seen to radiate, but nothing
further could be made out.
Occurrence.—The locality is given by Milne-Edwards simply
as Lake Tanganyika. Moore states* that he obtained this species
only off the west coast of the lake, and in nets and dredges worked
in water of about 20 fathoms. The experience of the third
Expedition shows that it occurs more widely distributed in the
lake, and may be taken in much shallower water than 20 fathoms.
Kasakalawe (south end): a single, rather young specimen,
taken under a stone about water-level, at the same time as speci-
mens of Potamon (Potamonautes) platynotus.
Mbete (south end): two or three young specimens came on
board my dhow clinging to the anchor-chain. The vessel was
probably anchored in two or three fathoms of water.
Among a number of specimens of Platythelphusa maculata,
dredged in 10-15 fathoms, principally at the south end of the
lake, are two or three small Crabs belonging to this species.
Vua, on the west coast: a single adult, seen from the dhow
crawling about in a foot or two of water, and caught in a hand-
net.
North end of the lake: a single specimen not full-grown.
* Op. cit. p. 280.
1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 271
PLATYTHELPHUSA MACULATA (Cunnington). (Plate XVII.
figs. 5 & 6.)
Limnothelphusa maculata Cunnington, Proc. Zool. Soc. 1899,
p. 698.
Limnothelphusa maculata Moore, ‘ The Tanganyika Problem ’
(London, 1903), p. 280.
Limnothelphusa maculata Rathbun, Nouv. Arch. Mus. Hist.
Nat. Paris, 4° sér., vil. (1905) p. 269.
As this species is now regarded as falling under the genus
Platythelphusa, in consequence of the examination of much more
extensive material, it becomes evident that it merits a more
complete specific description, which accordingly follows.
Description.—Carapace moderately convex, extending laterally
less than an orbit’s breadth beyond the external orbital angle.
Antero-lateral margins shorter than postero-lateral, with which
they are almost directly continuous. Number of spines on antero-
lateral margin variable, usually three, in addition to the spine at
the external orbital angle. Regions and sutures moderately
marked. Postero-lateral regions exhibit a series of small slightly
oblique granular ridges. Front more than one-third the width of
the carapace, margin perlated, and in extreme cases produced into
a small spine at each extremity. Sub-orbital spines little deve-
loped: no spine on the descending process of the front. Orbits
large, -24 width of carapace; orbital margins perlated. Eyes and
peduneles large. Chelipeds in the male unequal, subequal in the
female; merus rather short, trigonous, with distally a sharp spine
on the anterior margin ; carpus with two spines on inner margin
and one above the point of articulation with the hand. Fingers
of the larger chela gape considerably, are pointed and slightly
hooked, and carry a few larger crushing-teeth proximally, with
smaller teeth distally. Fingers of the small chela, and both chele
of the female (Plate XVII. fig. 5), distally meet closely in a sharp
cutting-edge, while proximally there are uniform teeth of moderate
size. Ambulatory legs of moderate length. Colour in life dark
brownish grey, with dark brown or reddish spots.
As now there has been obtained quite an extensive series of
specimens, some of which are larger than those examined before,
it seems worth while to furnish a further list ef dimensions, to
compare with those given under the original description of this
species. As in the case of P. armata, the larger chela of old
male specimens is of a great size, the length of hand and pollex
exceeding the total breadth of the carapace.
Adult male : mm.
ene thgoiscanapace mee \seeseny ees eee ere eens 12:9
iBveaddthpofrcarapacem ante cee oon eee 16-4
Hhgorango=ore owe Ml WCU copencossodoseeanooodoveconbacanuror 13:2
Ect v0 feito 1G yee VS 0 ac Mea he ere 6-1
Larger chela: greatest length of hand and pollex 16:9
Bs » greatest height of hand ............ 8:7
272 DR. W. A. CUNNINGTON ON THE BRACHYUROUS [Mar. 5,
Adult female (largest specimen) :
mm.
Taength of carapace © .........cecmeceueescoeenecemen cs 13°1
Brena tlaareCama Paces cacsegs-acine Sorevige arse ceataeeteee 16°9
Hirombo-or Oita wid hit caren -akee reer eeer ach ene tere: 13°2
WWadthvofsronk 261 atht-ereear fidabadopeaantinatetucs 6:1
Remarks.—While none of the specimens examined in 1899
carried ova, it is satisfactory, on comparison with this more ex-
tensive collection, to find that those regarded as adult were so in
reality. Female specimens of approximately the same size as that
described, but one of which is ovigerous, while the other carries
the already liberated young, occur in the recent collection.
Indeed we have further a female with total breadth of only
12:4 mm., but which is nevertheless ovigerous. The average
size of the eggs themselves, which are not quite round, 1s
IS) Se Ihss) iar,
The much smaller size of this species enables it to be distin-
guished at once from adult specimens of P. armata, but, apart
from that, the great relative breadth of the front and size of the
orbits are differences easily recognised. Among the number of
specimens which we now possess there exists considerable indi-
vidual variation in respect to the development of certain spines,
and this accounts for a slight discrepancy which may be noticed
between the foregoing description and that given in the first
instance. In the original type specimens, the spine on the carpus
of the cheliped, above the articulation with the hand, is indicated
but slightly, if at all. As it is, however, very well developed in a
number of individuals, it is now included among the specific
characters. Again, certain of the new specimens exhibit a small
spine at each extremity of the front, and have a slight indication
of sub-orbital spines, while others show no trace of these features.
A more complete description of the fingers of the chelipeds is
now given, and from my own observations I can add the colora-
tion during life.
Occurrence. —The type specimens described in 1899 were
obtained, according to the information supplied me by Mr, Moore,
from Kituta Bay (south end), while he had also taken specimens
at Niamkolo (south end) and Sumbu on the west coast. The
Crabs were said to have been captured in fairly deep water—never
less than 60, and from that to 500 feet deep. In his book on
Tanganyika, however, Moore states* that the specimens were
obtained in water varying in depth from 500 to 600 feet. He
also adds that the Crab occurs throughout the lake. With the
latter statement, the experience of the recent Expedition is quite
in accord; but whether the original individuals came from such
very deep water or not, it is certainly a fact that the species may
be found in much shallower regions.
Niamkolo Bay (south end): a large number of specimens
* Op. cit. p. 280.
1907.] ORUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 273
dredged among shells in about 10-15 fathoms. Also obtained in
rather shallower water, but seldom in less than about 8 fathoms.
Pembe (east coast): among shells dredged in about 10 fathoms.
Kirando (east coast), 1/12/04: dredged in about 10 fathoms.
Among these, a curiously blotched and not spotted individual.
PLATYTHELPHUSA CONCULCATA, sp. n. (Plate XVII. figs. 2 & 4.)
Description.—Carapace extremely flattened, extending laterally
less than an orbit’s breadth beyond the external orbital angle.
The whole body remarkably thin in appearance. Antero-lateral
margins shorter than postero-lateral, with which they are almost
directly continuous. ‘Three spines on antero-lateral margin, in
addition to that at the external orbitalangle. Regions and sutures
very ill-defined. Lateral regions exhibit a few small granulay
ridges. Front more than one-third the width of the carapace,
margin spinuliferous and with a sharp spine at each extremity.
Sub-orbital spines prominent; a small spine on the descending
process of the front. Orbits of moderate size, 18 width of cara-
pace; orbital margins perlated. Eyes and peduncles rather large.
Chelipeds in the male subequal ; merus trigonous, with distally a
sharp spine on the anterior margin, and a small blunt spine on
the ventral margin; carpus with two spines on inner margin, two
on outer margin, and one above the point of articulation with the
hand. Fingers distally meet closely in a sharp cutting-edge,
while proximally there are uniform teeth of moderate size.
Fingers of the larger chela (Plate XVII. fig. 4) have a few larger
teeth proximally. Ambulatory legs long and slender ; the anterior
margin of the merus, in each case, produced distally into two
sharp spines, of which the terminal one is inconspicuous on the
first and fourth legs. Colour (in spirit) yellowish brown, with
darker reddish-brown spots.
Dimensions as follows :-—
Male (probably adult) : fai
Meneth of carapace ..\sheass sn naae- 2-00 ne 10:0
Breadt ln olveaid PRE: ./Mas ce dastemal- oleae a> e9
Bvouto-oroibaly WiGtle i -l)jasety dae) eet ele ol
WW icitherotficonit 271), gue seas aaare tote. ade 50
Remarks.—It is unfortunate that we possess only a single
specimen of this species, which appears nevertheless to be quite
well marked. In size it is still smaller than P. maculata, while
the much flattened carapace and remarkable thinness of the body
(whence the name) at once arrest the attention. The great
development of spines is also striking, there being two additional
ones on the outer margin of the carpus of the cheliped and the
same number on the anterior margin of the merus of the ambula-
tory legs.
Occurrence.—-The only specimen obtained was associated with
274 DR. W. A. CUNNINGTON ON THE BRACHYUROUS _[ Mar. 5,
P. maculata, and dredged among shells in 10-15 fathoms of water
at the south end of the lake.
3. General Remarks.
The principal result of our extended knowledge of the Brachyura
of the African lakes is to make still more clear the very special
nature of the Crab-fauna of Tanganyika. As is found to be the
case in so many different groups of animals, the forms occurring
in Tanganyika are for the most part endemic, while those found
in the other big lakes are often of wide distribution. There occur
in Nyasa, as we have seen, three species of Potamonautes, one of
which, it is true, is described from that lake only. Inthe Victoria
Nyanza, we find Potamon (Parathelphusa) niloticus, a very widely
distributed form, and P. (Geothelphusa) emini, also known from
Abyssinia. From the Albert Edward Nyanza there comes also a
species of Geothelphusa. Thus, with the single exception of
P. (Potamonautes) orbitospinus, from Nyasa, the forms at present
known from these big lakes are by no means confined to them.
With Tanganyika it is quite otherwise. There are two species
of Potamonautes, one of whichis not known elsewhere, but beyond
this, three species of a unique and remarkable genus wholly re-
stricted to this lake. From the other lakes we have then only
representatives of the subgenera of Potamon—forms such as are
widely distributed in the tropical fresh-waters of the Old World.
From Tanganyika, while we have some representatives of these
normal African types, we have a preponderance of forms perfectly
distinct and occurring nowhere else. There is an indication, too,
of that richness of the Tanganyika fauna which is so noticeable
in some other animal groups. We know of five species of Crabs
from Tanganyika, three from Nyasa, and only two from the
Victoria Nyanza.
It is necessary to add a few remarks on the affinities of the
genus Platythelphusa. 'The species P. armata has been considered
to exhibit a distinctly marine appearance. Milne-Edwards stated
in the course of his original description that the Crab bore such a
resemblance to certain. marine or brackish-water Grapside that
we might relate it to that group, were it not for the development
of the abdomen and the absence of metamorphosis. In his book
on ‘The Tanganyika Problem,’ Moore* very rightly challenges
the value of such a character as the latter to the systematist. If
the absence of metamorphosis is the result of a particular habitat,
as we have reason to suppose, we ought not to take it into account
when we attempt to determine the aftinities of a newly-discovered
specimen. At the same time it would seem as if the resemblance
to the Grapside could be only a very superficial one, produced,
perhaps, by the more or less quadrilateral shape of the carapace.
* Op. cit. p. 236.
1907.] CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. 275
It must be admitted that the considerable development of
spines, which we find in all these species, is a feature more
commonly found among marine Crabs than among those inhabiting
fresh-water, and so would in itself convey a false impression on a
casual inspection. But we need not go outside the subfamily of
the Potamonine to find examples of Crabs which are just as
“‘marine looking,” as far as the development of spines is concerned.
This is true of a number of species of Parathelphusa, and when
we pass to the other subfamilies of the Potamonide, we have
Potamocarcinus, Valdivia, and Dilocarcinus, all extremely well
armed with spines. As with all these genera and subgenera,
there can be no doubt that Platythelphusa, despite its appearance,
finds its nearest allies in this group of forms which are essentially
and typically fresh-water in habitat.
It is a matter of more difficulty to decide which of the allied
forms are most closely related to Platythelphusa. The species now
known as P. maculata has been stated to be a primitive form™*,
on account of the little deflection of the front, the nature of the
antenne, and the spine-bearing margins of the carapace. These
features are equally characteristic of the other two species of the
genus, and do seem to show less specialisation than the deflexed
front and inflated gill-chambers of some of the semi-terrestrial
species of the subgenus Potamon. It is then not with the
latter, but near the genus Hydrothelphusa and the subgenus
Parathelphusa that we ought probably to place this distinctive
Tanganyikan genus.
The problem of the origin of Lake Tanganyika, about which so
much has been said and written, is of course intimately connected
with the questions which have just been discussed. If the lake be
the modified remains of part of an ancient ocean, we may expect
its inhabitants to show both a primitive character and a marine
aspect. These, Mr. Moore considers, are exhibited by all the
members of his “ halolimnic” group, among which are reckoned
the species of Platythelphusa. It has been shown, however, that
‘this genus has no better claims to a marine origin than other
representatives of the family, and at the same time, that while it
is not so specialised as certain allied forms which have adopted a
partially terrestrial mode of life, neither need it be considered as
the most primitive in the group. Still there is little evidence
that this form was ever anything but wholly aquatic, and it may
have become modified by the truly oceanic conditions prevailing
in Tanganyika, until it attained a superficial resemblance to
marine types.
In his report on the Macrurous Crustacea of the Expedition,
Dr. Calman shows} that they are to be regarded rather as
specialised than as primitive in character, and it may be asked
* Proc. Zool. Soc. 1899, p. 702.
+ Proc. Zool. Soc. 1906, p. 204.
276 CRUSTACEA OF THE THIRD TANGANYIKA EXPEDITION. [ Mar. 5,
why the same is not true of the Brachyura. An explanation is
really not far to seek. All the Macrura concerned are wholly
aquatic types, while among the Brachyura we have to institute
comparisons with forms which have partially accustomed them-
selves to a terrestrial existence. Such a profound change in
habits must produce an effect which, in comparison, would dwarf
any modification brought about within the limits of a single
medium.
There is gradually being accumulated a mass of information
concerning the other animal groups inhabiting Tanganyika, and
in nearly every case it is found that the forms are to a large
extent endemic and, moreover, very distinct and highly modified.
The explanation of this fact will be equally the explanation of
the remarkable character and variety of the Tanganyika Crab-
fauna, when compared with that of the other great African lakes.
There seems little doubt that it is to be found in a growing
divergence taking place in the lake during a prolonged period of
isolation.
EXPLANATION OF THE PLATES.
Prate XVI.
Fig. 1. Potamon (Potamonautes) orbitospinus (p. 259). Adult male, general view
from above. Nat. size.
Figs. 2-7. Series of figures of Platythelphusa and Potamon to show the frontal
region and illustrate the relations between the front, descending process,
and antenna (p. 267).
Fig.2. Platythelphusa maculata. X 5.
3. Platythelphusa coneulcata. X 5.
4. Platythelphusa armata. X13.
5. Potamon (Parathelphusa) niloticus. X 1s.
6. Potamon (Potamonautes) platynotus. X 2.
7. Potamon (Potamonautes) perlatus. X 2.
Prate XVII.
Fig. 1. Potamon (Potamonautes) platynotus (p. 264). Adult female, general view
from above. Nat. size.
2. Platythelphusa conculcata (p. 273). Male, general view from above. X 3.
3. Potamon (Potamonautes) platynotus (p. 264). Larger chela of adult female,
to show nature of dentation. X 1%.
4. Platythelphusa conculcata (p. 273). Larger chela of male, to show nature
of dentation. X 5.
5. Platythelphusa maculata (p. 271). Chela of female, to show nature of
dentation. xX 5.
6, Platythelphusa maculata (p. 271). Abdominal region of a female, from
below, to show the large size of the abdomen, and the degree to which
it covers the sternum. X 3.
Reference Letters.
ant. 1. Antennule. ex.orb. External orbital spine.
ant. 2. Antenna (second segment). s.so. Sub-orbital spine.
d.pr. Descending process of front. t.so. Sub-ocular tooth.
1907.] ON NEW SPECIES OF AFRICAN OLIGOCHZTE WORMS. 277
5). On Two new Species of the African Genus Microchetus
belonging to the Collection of Oligochzeta in the Museum
of Christiania. By Frank H. Bepparp, M.A., F.R.S.,
Prosector to the Society.
[Received February 1, 1907. |
(Text-figures 85 & 86.)
Dr. Robert Collett, the well-known chief official of the Christiania
Museum, was so good as to entrust me, some little time since,
with the collection of Oligocheta belonging to that Museum for
study and description. In examining the collection I found
three specimens, representing two species, of the Ethiopian genus
Microchetus* which I believe to be new to science, and of which
I beg to lay the following descriptions before the Society.
MICROCHATUS COLLETTI, sp. n.
I have the pleasure of dedicating this obviously new species to
Dr. Collett. The material consists of but one specimen, which is
entire, but considerably softened. It measures about 170 mm.
in length by 7 mm. in breadth after the clitellum ; the anterior
region of the body is wider. The colour is grey-brown, yellow
on the clitellum.
The sete have the usual paired arrangement found in this
genus, and commence upon the second segment of the body. The
sete are smaller upon the anterior segments and considerably
larger in the clitellar region, where they are quite twice the
length. The larger setz are ornamented. Some of the anterior
segments consist of two rings.
The clitellum (text-fig. 85) is very sharply marked off by its
colour and by the greater thickness of the body-wall in this
region. It commences with the xivth and ends with the xxivth
segment. The position of the clitellum is not anomalous for the
genus, and the specimen permits of no doubt upon the matter.
The clitellum is perhaps best described as ‘“ saddle-shaped” ; but,
as a matter of fact, the clitellar epithelium has also invaded the
ventral surface. There is, however, a diminution in thickness
indicated by an overhanging of the body-wall just ventralwards
of the lateral sete. This arrangement is interfered with in the
region of the genital papille by those structures, as will be seen
by the accompanying figure (text-fig. 85).
The nephridiopores are obvious and in front of the lateral pair
of sete.
The oviducal pores are very conspicuous upon the xivth segment.
Each lies between and in the same line with the ventral and
dorsal sete.
* In addition to the species known up to the end of last century (see Michaelsen,
Oligocheta in ‘Das Thierreich’), I. grisews has been subsequently described (see
Michaelsen, MT. Mus. Hamburg; xix. 1904),
278 MR. F. E, BEDDARD ON NEW SPECIES [ Mar. 5,
The genital papille (see text-fig. 85) are large and conspicuous ;
they occupy the xviith, xviiith, and xixth segments. They are
quadrangular in shape, with rounded angles, and each is continued
by a narrow ridge towards the middle ventral line. I traced the
sperm-ducts in the interior of the body as far as the segments
occupied by the genital papille, in connection with which I pre-
sume that they open. But I am unable to fix more precisely the
point of opening.
Text-fig. 85.
Microchetus colletti.
T have not found it altogether easy, in view of the condition
of the specimen and my unwillingness to injure it, as it is a
type, to ascertain the position of certain of the internal organs.
Assuming that the single pair of sperm-sacs is in the xth seg-
ment, the strongly marked gizzard lies in segment vil. This
segment is followed by two very thick septa, which thus separate
segments Vil. /viii. and viii. /ix. The dorsal vessel is double in parts,
and in segment ix. each half is much dilated, and a heart-like
structure is formed precisely like that which I first described in
Microchetus microchetus*, and which also occurs in other.species.
The last pair of lateral contractile hearts les in segment xi. The
calciferous glands are in segment 1x.
The spermathece, which are, as is the case with other species
of this genus, minute in size, open on to the boundary-line of
* Trans. Zool. foc. xii. p. 63.
1907. ] OF AFRICAN OLIGOCHATE WORMS. 279
segments x1./xil. and xii./xili. There are either three or four on
each side of the nerve-cord in each segment scattered between
the ventral region of the segment and the dorsal mid-line.
Copulatory glands are rather numerous in this species. There
are three pairs anteriorly, which le respectively in segments x.,
xi.,and xii. Each gland is double, as is commonly the case, being
composed of two sausage-shaped glands uniting to form a common
duct. They lie to the inside of the sperm-duct, which is almost
or quite in contact with them as it passes back to the external
pore. In addition to these three smaller pairs of glands there is
a single much larger pair lying two or three segments behind the
clitellum, and occupying a similar position in the body. ach of
these glands i is also double; but each tubular half is longer and
is coiled once upon itself. There are no conspicuous bundles of
genital sete associated with any of these copulatory glands.
The locality of the species is “ Zululand.”
MiIcrocH#TUS ZULUENSIS, Sp. Nn.
Two fragments contained in the same tube and from the same
locality as the species just described cleariy belong to the same
genus, but as plainly constitute another species of that genus
hitherto undescribed. Both fragments include the entire head
and body for a long way behind the clitellum. I do not give
exact measurements; the diameter of the anterior part of the
body is 12 mm. or so. The worms thus belong to a larger species
than Microchetus colletti.
One of the specimens (text-fig. 86) is more fully mature than
the other, and there is a difference in the number of the genital
papillee which is not attributable to immaturity. Iam thus able
to give a better account of external characters. But as the worms
are much softened my examination of the internal anatomy has
led to less satisfactory results. Nevertheless, I have been able to
make out certain anatomical facts which are of importance in the
discrimination of species.
The sete are so minute and difficult to see upon many segments
that I have found it impossible to map the regions of the body
by their aid only. Assuming, however, that the male pores are
upon the border-line of segments xiv./xv.,a very usual position for
them to occupy in this genus, and that the gizzard is in segment
vil., which is also the case with other species, I arrive at the
following determination of the position of the clitellum and of
other organs.
In the fully mature individual the clitellum, recognisable on
the xivth segment, is completely developed on the xvth, and
extends to the end of the xxviith. It is saddle-shaped, a bare
ventral area being left. In addition to the clitellum, segments
Xvi.-xxil. possess on either side a longitudinal band distinct from
the clitellum, though of the same appearance in general, coupled
with slight dissimilarity in colour, which is plainly the tubercula
280 ON NEW SPECIES OF AFRICAN OLIGOCHHTE WoRMS. [ Mar. 5,
pubertatis. This is confirmed by the fact that in the second im-
mature example without a clitellum the tubercula pubertatis were
nevertheless quite plain, though apparently occupying a segment
less.
Text-fig. 86.
Microchetus zuluensis.
The copulatory pupille (text-fig. 86) with their sete are very
obvious and very numerous in this species. In both specimens
there are two series of these papillee, one anterior to the clitellum
and one on some of the posterior segments of the clitellum. In
the fully mature individual with the completely developed clitellum
there is a single pair of these structures on segment x1., a papilla
of the pair being situated a little to the dorsal side of each
ventral pair of sete. These (the ventral) setee are also present on
the xith segment. These anterior papille are considerably larger
than those of the posterior series. The latter are in all five pairs
situated on segments xxiii.-xxvii. inclusive, thus immediately
continuing on the tubercula pubertatis. In the immature indi-
vidual there were the same papille upon the xith segment ; but in
1907.] THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 281
addition to these there was a smaller pair upon the xiith segment,
more ventrally placed, and in fact corresponding in position to
the posterior series. These latter papille in the present specimen
are fewer than in the adult, but they commence upon the same
segment, 7. é. the xxilird. There are four upon the right side of
the body and only two upon the left.
The gizzard is distinctly contained in two segments, which are
the vith and viith; the greater part of it, however, lies in seg-
ment vil. The mesenteries separating segments vi./ix. are thick.
The rather small calciferous glands are in segment 1x. ; the intes-
tine begins suddenly in segment xi. The dorsal blood-vessel is
dilated in segment ix. I imagine that this species is one of those
which only possess one set of testes, funnels, &e. For the sperm-
sacs and reservoirs consist of one pair in the ixth segment attached
to the posterior wall of that segment and a pair in the xth
attached to its anterior wall, 7. ¢. the same septum as that which
bears the sperm-sacs. The two sacs of each side of the body seem
to communicate, and I take the anterior pair to be the sperm-sacs
and the posterior pair to contain the funnels.
T was quite unable to find any spermathece. 1 am unwilling,
however, to assert that these organs are absent. If they happened
to contain no sperm their minute size and the softened condition
of the worm would render it at least very difficult to detect them.
The only other species, as it appears, in which no spermathecz
have been detected is Dr. Michaelsen’srecently described I. grisews*.
But as this latter species is holandrous it cannot be confused with
M. zuluensis, than which it is also a good deal smaller,
March 19, 1907.
Dr. Henry Woopwarp, F.R.S., Vice-President, in the Chair.
The Secretary read the following report on the additions that
had been made to the Society’s Menagerie in February 1907 :—
The registered additions to the Society’s Menagerie during the
month of February were 76 in number. Of these 27 were
acquired by presentation and 10 by purchase, 36 were received
on deposit, 1 by exchange, and 2 were born in the Gardens.
The total number of departures during the same period, by death
and removals, was 188.
Among the additions special attention may be directed to :—
A Long-tailed Goral (Wemorhedus caudatus), from Korea, new
to the Collection, presented by Mr. C. F.G. Bilbrough on Feb. 5th.
A Harpy Hagle (Thrasaétus harpyia) from South America,
purchased on Feb. 8th.
Myr. Herbert F. Standing read a paper, illustrated by lantern-
* Loc. cit. (on p. 277).
282 ON NEW ANIMAL PARASITES. [ Mar. 19,
slides and large series of photographs and specimens, on recently
discovered subfossil Prosimiz from Madagascar, in which he dis-
cussed their affinities with extant Lemurs and with the higher
Primates. The remains were obtained in the muddy bed of a swamp
formed by the blocking-up of the river Mazy by a lava-flow, at from
a few inches to 3 or 4 feet below the surface. They consisted of
a large number of skulls and limb-bones of Lemurs and Lemur-
like animals. This great amount of material enabled the author
to corroborate the view, previously put forward by Dr. Forsyth
Major, that the extinct Lemurs of Madagascar were, in many
respects, intermediate between existing Lemurs and Monkeys,
and to express his belief that the New World Monkeys and the
Lemuride, as well as the Malagasy Indrisine, had a common origin.
He also stated his opinion that, in view of the recent additions to
our knowledge of the Prosimiz and of what the present collection
revealed with regard to their close relationship to the Apes, it
was not possible to separate the Primates, as hitherto, into the
two suborders Lemuroidea and Anthropoidea.
This paper will be printed entire in the ‘Transactions,’
The following papers were read :—
1. Descriptions of some New Species of Animal Parasites.
By L. W. Sampon, M.D., F.Z.S.*
[Received March 19, 1907. }
WELLCOMIA MITCHELLI Sambon.
Abstr. P. Z. 8. 1907, p. 15 (March 26).
Habitat. Small intestine of Pedetes cafer.
Only females found, 12-15 mm. long and about 1 mm. broad.
Characterised by the presence of a conical ovipositor 2-3 mm.
long, placed ventrally on the anterior third of the body, 2-
3°5 mm. from the cephalic extremity, and by a spirally twisted
tail, 2-3 mm. long, terminating in a fine point. Body semitrans-
parent. Head tapering anteriorly. Mouth trilabiate ; cesophagus
long and terminating in a spherical bulb. Anus open ventrally
at 3-4 mm. from tail-end. Eggs smooth, oblong, asymmetrical,
and measuring 60-65 p by 28-32 p.
SPARGANUM BAXTERI Sambon.
Abstr. P. Z. 8. 1907, p. 16 (March 26).
Habitat. Connective tissue of Man.
Long, flat, unsegmented body, 15 em. long and 1:5 mm. broad,
* [The complete account of the new species described in this communication
appears here; but since the names and preliminary diagnoses were published in the
« Abstract,’ the species are distinguished by the name being underlined.—Ep1ror. |
ESO) ON NEW HEMOGREGARINES. 283
with numerous irregular transverse folds and a distinct longitu-
dinal groove on ventral surface. Anterior extremity 2-5 mm.
broad; head completely invaginated. Posterior extremity 2 mm.
broad, with shallow median slit. Extracted from an abscess on
the thigh of a Masai in British Central Africa.
ScHISTOSOMUM MANSONI Sambon.
Abstr. P. Z. 8. 1907, p. 16 (March 26).
Habitat. Blood-vessels of Man.
In the Congo Free State, in other parts of Africa, and in the
West Indies there is a form of Bilharziasis clinically and patho-
logically similar to the Asiatic form caused by Schistosomum
japonicum, and unlike the classic Kast African form due to
S. hematobium. The eggs of the species which causes this peculiar
form are never found in the urine, but seem to be eliminated
through the intestine only. They differ from those of S. hema-
tobiuwm in having a broad lateral spine totally different in size,
shape, and position from the small, straight, terminal spe which
characterises the ova of S. hematobiwm. Hitherto, the laterally
spined ova, usually observed in Egypt in cases of mixed infection,
have been looked upon as having been distorted while passing
through the rectal mucosa. Sir Patrick Manson suggested several
years ago, that the laterally spined ova found in the feces of
patients, and never in the urine, might represent a new species.
In appreciation of this, one of his many genial intuitions, the new
species 1s dedicated to him.
2. Descriptions of five New Species of Heemogregarines from
Snakes. By L. W. Samson, M.D., F.Z.8S., and C. G.
SeLicmMann,. M.D., F.Z.8.*
[ Received March 19, 1907. |
H.&®MOGREGARINA POCOCKT Sambon.
"Abstr. P. Z. 8. 1907, p. 16 (March 26).
Habitat. Erythrocytes of Indian Python, Python molurus L.
Club-shaped, 14-16 p long. Anterior extremity rounded, 3-15
broad. Posterior extremity attenuated and recurved. Cytoplasm
more or less granular. Nucleus median or nearer posterior ex-
tremity, large, oval, and with coarse, deeply staining chromatin
granules. Parasite lies parallel or obliquely to long axis of host-
cell, of which it occupies about two-thirds, without causing much
alteration beyond displacement of nucleus.
* [The complete account of the new species described in this communication
appears here; but since the names and preliminary diagnoses were published in the
* Abstract,’ the species are distinguished by the name being underlined.—Eprrokr. |
284 ON NEW HH MOGREGARINES. [ Mar. 19,
H#MOGREGARINA SHATTOCKI Sambon.
Abstr. P. Z. 8. 1907, p. 17 (March 26).
Habitat. Erythrocytes of Diamond Snake, Python spilotes
(Lacép.).
Club-shaped. Some forms more slender, 14-15 long and 2 yu
broad, with both extremities rounded and differing only slightly
in thickness. Other forms more pay somewhat similar to those
of Python molurus, but larger, 22 uw by 4. Nucleus median
and very large, 9p by 4p. Host-cell sometimes slightly distorted,
nucleus pushed to the periphery.
H 4M0GREGARINA REFRINGENS Sambon.
Abstr. P. Z. S. 1907, p. 17 (March 26).
Habitat. Erythrocytes of Hoary Snake, Pseudaspis cana L.
Crescentic, bean-shaped, and discoidal forms occur. The slender
crescentic forms have a long, oval, and more or less central nucleus.
The bean-shaped forms measure 10-12 y in length by 5-6 « in
width; they have a wide central nucleus, and their cytoplasm is
literally crammed with rounded, highly refractive granules. Host-
cell unaltered beyond occasional displacement of nucleus.
H £M0GREGARINA MANSONI Sambon.
Abstr, P. Z. 8. 1907, p. 17 (March 26).
Habitat. Erythrocytes of Testaceous Snake, Zamenis flagelli-
Forms L.
Oval or bean-shaped cyst 12-13 » long by 5-6 w broad, en-
closing club-shaped parasite doubled up in the form of a letter U
with both branches of equal length and closely applied. Nucleus
median and situated near bend at one pole of cyst. Chromatin
arranged in transverse parallel lines or in concentric circles.
A characteristic feature is the almost constant presence of two
large chromatoid granules usually placed one on each side of
nucleus. Host-cell unaltered, nucleus slightly displaced.
H £MOGREGARINA RAREFACIENS Sambon.
Abstr. PB. Z. 5. 1907, p. 17 (March: 26):
Habitat. Erythrocytes and leucocytes of Couper’s Snake,
Coluber corais var. couperi Holbr.
1. Slender, elongate, cylindrical forms 14 p long by 1:5 p broad,
sometimes presenting a refringent granule or vacuole at each
extremity. Host-cell apparently unaltered or only slightly
Pear trophied.
Large bean-shaped forms 12-13 pw long by 4-5 pw broad, with
Res more or less granular and occasionally vacuolated.
Nucleus small, round, median! with fine chromatin grains. The
host-cells containing ‘this form measure about four times the
normal size, and are entirely dehzemoglobinized and greatly atten-
uated. Their nucleus is hypertrophied. Sometimes two or even
three parasites may be found in the same host-cell.
1907. | ON MAMMALS FROM COGUNO, INHAMBANE. 285
3, The Rudd Exploration of South Africa.—VII. List of
Mammals obtained by Mr. Grant at Coguno, Inham-
bane. By OLp¥FieLp THomas, F.R.S., F.Z.8., and R. ©.
Wrovuenton, F.Z.8.
[Received February 21, 1907. ]
No part of South Africa was more inadequately represented in
our National Museum than the Portuguese territories north-east
of Delagoa Bay, and we are glad to say that the fine collection
from Inhambane of which we now give an account does much
towards filling up the lacune. Mr. Grant found a good collecting
place at Coguno, about 75 miles to the south-west of the town of
Inhambane, and obtained there the whole of the present series,
which numbers. 212 specimens, belonging to 59 species.
Of these we have described 6 as new, several of them being
particularly striking forms, notably a Galago and a Petrodromus.
And besides the novelties, specimens that are of very special
value are the topotypes of such of Peters’s species as were described
from Inhambane, these being the first of a series of Petersian
topotypes which we hope Mr. Grant may in time be able to com-
plete, since all S.E. African Mammalogy seems to circle round the
species obtained during the epoch-making voyage of the famous
German zoologist, and but little satisfactory work can be done until
good modern specimens of all his species are available for study.
As before the whole of this valuable collection is presented to
the National Museum by Mr. C. D. Rudd.
Mr. Grant’s notes on the Inhambane District are as follows :—
“Tnland from Inhambane the country is composed of more or
less undulating, sandy flats, densely bushed and timbered ; often
for many miles the only open spaces are the clearings made by the
natives for their kraals and cultivated lands.
“Some of the river-valleys are more open, consisting of patches
and stretches of forest alternating with open plains, generally
thickly covered with palmetto.
“The rivers, excepting the Intanime, are merely huge, dense
reed-beds, through which the actual watercourse is often traceable
only with difficulty.
“The natives are very numerous and are mostly Mchopee, with
a small sprinkling of Machangaan, and all are great hunters and
trappers, and gave considerable help throughout the trip, either
by giving information as regards species or in bringing in
pean
‘My head camp lay close to the Commando of Coguno, which
is about 130 kilometres by the Panda Road from Inhambane ; :
thence short trips and excursions were made into the surrounding
country as necessity demanded.
“Owing to the denseness of the bush and the comparative
flatness of the veldt local guides were always necessary.
Proc. Zoou. Soc.—1907, No. XX. 20
286 MESSRS, 0. THOMAS AND R. C. WROUGHTON ON | Mar. 19,
‘“¢So far as I could learn, the District has never been surveyed,
so that the altitude of Coguno, or other places, is unknown,
but probably no part exceeds 1000 ft. The actual position of
rivers and localities can only be roughly estimated.
“Throughout my stay the climate was delightful ; the average
temperature being around 80° in the shade, occasionally reaching
88°; it being the dry season, there was little rain.
“The following species were not obtained but exist at Coguno,
and the native names may be of interest. They are:
COL LAN HAG. MaRS A oe HER OO Geo CES Shalu.
WING OS ORGCUIS peer eres ee: Lengo.
Thryonomys swinderenianus ... Shleti.,.—C. H. B. G.
1. Papro porcArius Bodd.
Flat skin.
“‘ Native name, ‘ Infeni.’
“ This species was not seen in the neighbourhood of Coguno
Camp, but is said to be common to the North and West. The
flat skin sent was obtained from a native who killed it some
20 miles to the northward of my camp.”—C. H. B. G.
2. GERCOPITHECUS PYGERYTHRUS ERYTHRARCHUS Peters.
Bio WOR WHO Qe WAGs, WE.
Mr. Pocock informs us that he considers Peters’s erythrarchus,
of which these are topotypes, should be ranked as a subspecies of
pygerythrus F. Cuvy.
“ Native name, ‘ Makaku’ or ‘ Ihow.’
“¢Gommon and inhabiting the denser forests where it lives on
the wild fruit and berries. Never observed in large troops ; single
old males were sometimes seen, but were usually unapproachable.
They are at all times wary and extremely difficult to secure.
Cercopithecus albigularis exists in the district, but is very rare
and local and even the natives were unable to obtain me speci-
mens; it is known to them as ‘ Glanglanu ’.”C. H. B. G.
3. GALAGO GRANTI, sp. n.
3. 1617, 1618, 1619, 1660, 1662. Q. 1517, 1653, 1663.
A member of the moholt group, with an unusually bushy black-
tipped tail.
Fur fine and soft, about 15 mm. in length on the back. General
colour above drab-brown, rather darker than Ridgway’s “ drab.”
Under surface cream-buff, the basal three-fifths of the hairs slaty.
Light nose-line and black orbital patches as in G. moholi. Outer
side of fore limbs light drab, lightening to white on the hands ;
inner side like belly. Hind limbs as usual dull cream-buff
throughout, rather duller than on the belly. Tail long, unusually
bushy, the hairs attaining a length of 20-25 mm., those of true
moholi but little more than half this length ; in colour the basal
three-fifths of the tail is drab-brown like the body, gradually
darkening terminally to blackish brown,
1907. | MAMMALS FROM COGUNO, INHAMBANE. 287
Skull readily distinguishable from that of G. moholi by its much
longer muzzle, the palate-length being over 17 mm., as com-
pared with 15 in the type of moholi, and all the other muzzle
measurements in proportion. The bulle also are lower and
less abruptly swollen anteriorly, and the canines appear to be
rather heavier.
Dimensions of the type :—
Head and body 158 mm.; tail 237 ; hind foot 63; ear 43.
Skull—greatest length 45 mm.; basal length 35; zygomatic
breadth 28; mastoid breadth 23°5; tip of nasals to back of
orbits 25; palate-length 18; front of canine to back of m® 15-6.
Type. Adult male. B.M.no.6.11.8.7. Original number 1662.
Collected 26 August, 1906.
This Galago differs from G'. moholi by its long bushy black-
tipped tail and by its markedly longer muzzle. Peters’s G. mos-
sambicus, the skull of which is figured by him under the name of
senegalensis, has a short muzzle as in the true moholi.
With regard to G. conspicillatus I. Geoft., Prof. Trouessart
kindly informs us that the tail of the type is only 20 mm. in total
breadth, therefore exactly as in moholi, not as in the Inhambane
species.
We have had great pleasure in naming this interesting little
‘‘ Bush-baby ” after its captor, Mr. C. H. B. Grant, who has
collected the whole of the immense number of mammals recorded in
the present series of papers, and has thus ably utilised the splendid
opportunity afforded him by the generosity of Mr. Rudd.
“ Native name, ‘Suwanjati.’
‘¢Common and inhabiting the forests. It is strictly nocturnal,
sleeping during the day in the hollow trees, where it may generally
be taken in small family parties. ‘This species like many others
is eaten by the natives.
“ G. crassicaudatus was several times heard calling in these
parts at night, and I saw one skin worn by a boy; none, however,
- could be secured by myself or the natives, although I offered good
rewards fora specimen. It is known to the natives as ‘ Gerile ’.” —
(Op Jal, 16) (Cr
4, SCOTOPHILUS NIGRITA Schreb.
$. 1577, 1578, 1621,1656. 9. 1579, 1616, 1665, 1671, 1672,
1673.
In this series there would seem to be two forms distinguished
by size. The larger corresponding closely to, if not identical with,
the specimens from Klein Letaba provisionally identified by
Thomas & Schwann (P. Z. 8. 1906, p. 577) as migrita. The
presence in one place of two forms differing in size seems to be a
common occurrence in this group, as for instance, planirostris and
viridis of Peters from the Zambesi Valley, and nigrita herero
and damarensis of Thomas from 8. W. Africa. We are of opinion
that it is safer to include all these specimens provisionally under
nigrita until the group can be worked out in its entirety.
20%
288 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 19,
“‘ Native name, ‘ Mongavilane.’
“Quite the commonest of the Bats at Coguno. Generally
appears long before dark, flying fairly high, and can be easily
secured with a shot-gun. Numbers 1621 and 1673 appeared to
me different from the others, from which they were easily
distinguishable on the wing.”—C. H. B. G.
5. ScoTOPHILUS SCHLIEFFENI Peters.
Go UDO, Gaile
This is the most southern locality from which this small Scoto-
philus has been recorded, the next being Marungu, whence
Dr. Noack described his S. minimus.
6. GLAUCONYCTERIS PAPILIO Thos.*
¢. 1580, 1603. ©. 1620.
Additional specimens of the rare and remarkable Butterfly-
bats of the genus Glawconycteris are extremely welcome. The
present examples agree in size and the colour of their heads with
G. papilio, and equally differ from the white-headed and white-
bellied G. variegatus Tomes, of Damara-land.
“‘ Native name, ‘ Mongavilane.’
“ Apparently uncommon, as the three specimens taken were
the only ones observed. They appear about the same time as
Scotophilus, but have a more butterfly-like flight, which easily
distinguishes the species at a good distance.” —C. H. B. G.
7. PIPISTRELLUS NANUS Peters.
©. 1668.
Quite like the specimens from Legogot mentioned in the last
paper on the Rudd Collections (Thomas & Schwann, P. Z. Sh
1906, p. 780).
“ Native name, ‘ Mongavilane.’
“These little Bats were fairly common, but, owing to their
small size and rather late appearance, a big series was not
obtainable.”—C. H. B. G.
8. AMBLYSOMUS OBTUSIROSTRIS Peters.
B Ws, IGS7, WIM, De Webs)
“Caught in run in thick bush.”
These valuable specimens are practically topotypes of Peters’s
species, which was described from the neighbourhood of Inhambane
town.
Their bellies are of a peculiar coppery brown, and the lighter
basal portion of their dorsal hairs is also strongly tinged with the
same colour. In A. chrysillus Thos. & Schw., from Delagoa Bay
the same parts are white or yellowish white. The hairs of the
back are about 5 mm. long in 4. obtusirosiris, 6-5 mm. in
A. chrysillus.
* Ann. & Mags. N. Hi. (7) xv. p. 77, 1905.
1907. ] MAMMALS FROM COGUNO, INHAMBANE. 289
“* Native name, ‘ Tukunve.’
“ Fairly common and forming runs just below the surface in
the sandy ground, in the forest and bush. Owing to the peculiar
nature and smallness of the runs it was found impossible to trap
it and it could only be taken when seen working.”—C. H. B. G.
9. PETRODROMUS SCHWANNI, sp. n.
6. 1521, 1523, 1536 1539, 1557. 9. 1500, 1515, 1522, 1524,
1537, 1538, 1552, 1553, 1554, 1555, 1556.
Allied to P. sultan* Thos., but skull shorter and the tail more
hairy and more finely scaled.
General characters as in P. sultan, the peculiar round-headed
bristles underneath the tail quite as in that species. Colour
sunilar throughout, except that the dull russet-brown dorsal area
is broader and more diffused, extending nearly across the back,
instead of forming a comparatively narrow line down the spine
sharply separated from the grey of the shoulders and flanks.
Face-markings, limbs, and under surface all quite as in sultan.
Tail rather longer than in suléan, and the scaling finer, 13 rings
of scales to the centimetre instead of 10 as in sultan; upper
surface well haired throughout, the scales nearly or quite
hidden—(in P. sultan the upper side of the tail is practically
naked and the large scales are clearly visible). In colour the
upper surface is deep black throughout, and the lower dull buffy,
not sharply contrasted; extreme base below pale flesh-colour.
Long knob-headed bristles black throughout.
Skull markedly shorter than in P. swltan, but the shortening is
chiefly in the muzzle, the brain-case being of about the same size.
Hinder edge of nasals level with the front of the anteorbital rim,
and falling about one millimetre short of the frontal processes of
the maxille. In P. sultan the nasals are of about the same
length, but end further forward. P* (the fourth tooth from the
back) comparatively small and narrow.
Dimensions of the type (measured in the flesh) :—
Head and body 192 mm.; tail 181; hind foot 57; ear 36.
Skull—greatest length 53 mm.; basal length 47; greatest
breadth 29°5; length of nasals 20; interorbital breadth 9;
breadth of brain-case 20; length of upper tooth-series 27; front
of p’ to back of m* 11°5.
Type. Old male. B.M. no. 6.11.8.32. Original number 1557.
Collected 5 July, 1906.
The occurrence of this handsome Petrodromus in Inhambane
was already recorded by W. L. Sclater?, who had received a
specimen collected there by Mr. H. F. Francis. With only a
single specimen, however, he did not think himself justified in
distinguishing it from P. sultan, and it is therefore placed in the
‘Mammals of South Africa’ under the latter name.
* P.Z.S. 1897, p. 435. Originally published as P. suJtani by a printer's error,
but the mistake was corrected P. Z.S. 1897, p. 928.
+ Mainm. S. Afr. 11. p. 155, 1901.
290 MESSRS. 0. THOMAS AND R. ©. WROUGHTON ON [Mar. 19,
In Herr O. Neumann’s revision of the genus* the primary
division of the species is made by the hairiness or nakedness of
the upper side of the tail, but the occurrence of this species,
obviously a close ally of P. sultan but with a hairy tail, shows that
this character is of but little importance. We should rather
divide the species, as in Thomas’s original paper, by the structure
of the caudal bristles, in which respect P. schwanni agrees with
P. sultan alone of all the described forms.
No Petrodromi of this type have been as yet recorded between
Inhambane and the mainland opposite Zanzibar, a distance of
some 1200 miles.
Weare glad of the opportunity of linking with this fine species
the name of Mr. Harold Schwann, who has hitherto been so
closely connected with the working out of the Rudd Collection.
‘“‘ Native name, ‘ Nyakole.’
“Very common and inhabiting only the dense thickets, where
it has regular runs, in which it is easily trapped. It has all the
actions of the Macroscelidide, carrying the tail almost perpen-
dicular when running. Exclusively diurnal and insectivorous.”—
CoH BG:
10. Crocipura sytv1A Thos. & Schw.
6. Ldai3:
“ Native name, ‘ Nongi’ (without distinguishing species).
‘* Not common and confined to the reed-beds and swamps along
the river-valleys, especially the Inyasuni.’—C. H. B.G.
11. CroctpURA FLAVESCENS FLAVIDULA Thos. & Schw.
Ge V5e2 1574:
12. Fevis ocrEATA Gmel.
2. 1636.
We hesitate to identify this single specimen with any particular
subspecies until the arrival of further specimens from Portuguese
Kast Africa.
““ Native name, ‘ Igoye’ or ‘Simange.’
‘“‘ Apparently very uncommon, as the specimen obtained was
the only one taken or observed.”—C. H. B. G.
13. VIVERRA CIVETTA Schreb.
OP One
“‘ Native name, ‘ Fungwi.’
“‘ According to the natives common, although I did not observe
more than the one specimen, except a couple of skins in the
possession of natives. Apparently inhabits the thickest of
forests.” —C. H. B. G.
* Zool. Jahrb., Syst. xiii. p. 541, 1900.
1907.] MAMMALS FROM COGUNO, INHAMBANE. 291
14. GENETTA sp.
3g. 1566. @. 1562.
Allied to G. letabe Thos. & Schw.
‘“‘ Native name, ‘Simba.’
** Not very common and found everywhere, especially near
native habitations. This species, with all the cats, is eaten by
the natives.”—C. H. B. G.
15. Muncos GALERA Erxleb.
3. 1645.
“* Native name, ‘ Shikoko.’
“« Apparently common and generally living in the reed-beds and
swamps; the specimen sent, however, was caught some little
distance from water and near a Kaftir kraal, where it was
probably on the prowl after a stray chicken. Nocturnal only.”—
C. H. B.G.
16. Muneos (IcHNEUMIA) GRANDIS Thos.
3. 1535.
Whether this animal, the external characters of which are quite
as in southern examples of MW. albicauda, should be treated as one
of the subspecies of that form, or whether several of these should
be raised to specific rank, is a question which cannot be settled
without further material. Immature specimens would be of
particular value as the characters rest mainly on the structure
of the teeth, which get worn down in adult life.
‘‘ Native name, ‘Sanganye.’
“By no means common and generally found near habitations,
which they visit for the chickens, &c.”—C. H. B.G.
17. MunGos CAFER Gm.
“Native name, ‘ Shlaushlwa.’
_ The flat skin sent was the only one seen during the trip. It
is evidently rare, as many of the natives did not know its name
and some even had never before seen one. The boy from whom
I took it said he caught it in the reedy bed of the Inyamatanda
River.”—C. H. B. G.
18. CROSSARCHUS FASCIATUS SENESCENS, subsp. n.
3. 1558, 1628, 1654.
“Shot in thick bush.”—C. H. B. G.
Much greyer than the true fasciatus.
Size rather larger than in Zululand examples of fasciatus,
approaching that of the Ruwenzori form, C. f. macrurus Thos.,
but the tail not lengthened. General colour conspicuously greyer
than in fasciatus, the nape, fore-back and flanks of a clear
cinereous grey, entirely without fulvous suffusion. The stripes
normal in number and position, but the light ones white along
992 MESSRS. O. THOMAS AND R. C. WROUGHTON ON Mar. 19
?
their anterior edge, only becoming a little fulvous posteriorly ;
in fasciatus each light stripe 1s fulvous anter iorly becoming darker
fulvous poster iorly. Head finely grizzled grey, blacker on the top
of the muzzle. Ears grey, without fulvous suffusion. Limbs
grizzled grey, dar kening terminally on the hands and feet to
black.
Skull of normal proportions, but the teeth unusually large.
Dimensions of the type (measured in the flesh) :—
Head and body 364 mm.; tail 236; hind foot 73; ear 24.
Skull—condylo-basal length 73 mm.; basal length 68; zygo-
matic breadth 39; palate- length 39; greatest diameter of p 74.
Type. Adult male. B.M. no. 6. 11.8.51. Or igimal number
1628. Collected 13 August, 1906.
This beautiful grey form of the Striped Mungoose may be
readily distinguished by the reduction of the fulvous suffusion of
the fur, this being only present along the posterior bor der of each
dorsal light band, while in the orher forms some trace of it
oceurs all over the body; the clear grey nape and shoulders of
C. f. senescens are especially noticeable.
The young specimen no. 1558 is more like the typical form.
‘“‘ Native name, ‘ Gale.’
‘“* Apparently fairly common, but dithcult to secure owing
partly to its wariness and partly to its often inhabiting the
denser parts of the bush. The specimens obtained were shot
during native hunts and were taken from troops of perhaps eight
individuals. It is distinctly gregarious and diurnal in habits,
living principally on coleopterous insects and the eggs and young
of ground-breeding birds.” —C. H. B. G.
19. MELLIVORA RATEL Sparrm.
‘“‘Native name, ‘ Sididi.’ ”—C. H. B. G.
20. Icronyx CAPENSIS Kaup.
1534, 1655. 9. 1575
‘“‘ Native name, ‘Shingemani.’
“Fairly common everywhere, especially near native kraals,
where it makes itself a considerable nuisance by stealing chickens.
Nocturnal only.”—C. H. B. G.
21. FUNISCIURUS SPONSUS, sp. 1.
3. 1596, 1600, 1608 (yg.), 1609, 1610. 9. 1547, 1560, 1561
1611, 161 2.
A Funisciurus intermediate in size between cepapi and palliatus,
with the colour pattern of the former and the bright colouring of
the latter.
Size rather smaller than in /’. palliatus.
Fur of back soft, 12 mm. long; that of tail 30-40 mm.
General colour above erey-br own freely grizzled with very pale
buff or yellow; the individual hairs black or dark brown, each
1907. ] MAMMALS FROM COGUNO, INHAMBANE. 293
with two rings (one near base and one subterminal) yellowish
white ; the belly orange-red, the hairs unicoloured to their bases.
Head igen the forehead backwards coloured the same as the back ;
face below the eyes, chin and throat orange like the belly. Outer
side of limbs coloured like the back, but a strong rufous tinge on
thighs; inner side orange like the belly. Tail coloured like the
back but with a strong rufous tinge at the sides and below; the
individual hairs with two pale rings, each 5 mm. wide, separating
three black rings of same width, with a long (10-12 mm.) ferru-
ginous tip.
Skull as in palliatus but decidedly smaller.
The following are measurements of the type (which scarcely
differ from those of others of the series). For convenience of
comparison corresponding measurements of a normal adult speci-
men of F, palliatus ornatus Gray from Zululand are added between
brackets.
Head and body 197 (200) mm.; tail (cire.) 195 (218); hind foot
41 (45); car 20 (18).
Skull—greatest length 48 (52) mm.; basilar length 37 (40);
zygomatic breadth 28 (30); interorbital breadth 15°5 ml 5); nasals
13°7 (15°5); upper molar series 8°7 (9°5); bulle 10°5 (11).
Type. Adult female. B.M. no. 6.11.8.63. Original number
1560. Collected 5 July, 1906.
Through the courtesy of Dr. Péringuey we have been able to
examine a Squirrel from St. Lucia Bay, Zululand (South African
Museum, no. 4361), assigned by Mr. W. Sclater * to frerei Gray,
under the impression that the type of the latter was obtained by
Sir Bartle Frere in Natal. But the typical specimen was received
by the British Museum in 1873, about which date Sir B. Frere
veturned to England from Zanzibar, and he did not go to South
Africa till some years later. The type locality “‘ Zanzibar” given
by Gray for his “ Macroxus anumulatus frere” is therefore
undoubtedly correct. The Zululand specimen, though paler in
colour, agrees in all essential characters with the present species,
to which we have no hesitation in assigning it. The range
of sponsus is therefore from Inhambane to Zululand, where it
coexists with F. palliatus ornatus Gray, and this fact and its
smaller size ee justify us in classing it as a distinct species.
‘““ Native name, ‘ Shintsi.’
““ Extremely common and found everywhere in the forests and
thickets, especially near native clearings; as many as half a
dozen can be seen at one time running about in the trees. The
alarm call is a bird-like chatter. It is taken in vast numbers by
the natives, with whom it is an especial delicacy.” —C. H. B. G.
As we have had occasion to refer to the true locality of frerei,
we further take this opportunity of recording that, allowing for
the fact that it is a young individual, we find on comparison that
the type of frerci agrees quite closely with the specimens assigned
* Mamma. S. Afr. ii. p. 7 (footnote), 1901.
294 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON’ [ Mar. 19,
by Herr Neumann to his /. palliatus suahelicus from the same
region, and we have no doubt as to their identity; swahelicus
Neum. must therefore give way to the older frerei Gray.
22. TATERA LOBENGULH BECHUANZ Wrought.
3. 1484, 1488, 1491, 1499, 1531,1532. 9. 1483, 1487, 1489,
1494, 1514, 1533.
These specimens, though less pale than typical bechuane from
the Bechuana Desert, are very close to that form. Their receipt
from Inhambane is interesting from the point of view of the
distribution of the species. The type of lobengule was described
from Matabililand and local races were subsequently named from
Kuruman, Matopo, and Salisbury, all places which may bedescribed
as on the left bank of the Limpopo. In the collection from the
Zoutpansberg District on the right bank, though more than one
species of 7atera were found, there was no specimen referable to
lobengule. More recently, however, in a small coljection from
Pietersberg District on the right bank of the Limpopo but much
lower down than Zoutpansberg, some specimens scarcely distin-
guishable from the present series were found. Broadly, however,
the area between the Limpopo and Zambesi Rivers would seem to
be the home of the species, which takes quite well-marked local
forms in various parts of its habitat.
‘“‘ Native name, ‘ Singaan.’
“ Quite the commonest rat in the district and found everywhere,
especially in and around the native clearings.”—C. H. B. G.
23. OTOMYS TRRORATUS CUPREUS Wrought.
3. 1583, 1584, 1586. 2. 1585, 1587, 1588.
These specimens are all young but agree closely with O. irroratus
cupreus from Zoutpansberg.
‘“¢ Native name, ‘ Woti.’
“Not common and only taken in the reed-bed and swamps
along the river.”—C. H. B. G.
24. ARVICANTHIS DORSALIS A. Sm.
do. 1498, 1519, 1614, 1625, 1626, 1630, 1634,1647. 2. 1486,
1607, 1615, 1623, 1633, 1638.
“ Native name, ‘ Maklangane.’
“Tt is here far commoner than in any locality where I have
previously taken it. Found everywhere, both in the forest and
bush and the native lands, where it has regular tracks and runs,
sunilar toits congener at the Klein Letaba. Exclusively diurnal.”
—C€. H. B. G.
25. SACCOSTOMUS CAMPESTRIS Peters.
3. 1495, 1505, 1506, 1508, 1509, 1597, 1606, 1629, 1639.
Q@. 1479, 1481, 1482, 1490, 1501.
These specimens are practically topotypes of S. fwscus Peters,
1907. ] MAMMALS FROM COGUNO, INHAMBANE. 295
described from Inhambane, but as they are indistinguishable from
S. campestris, it seems probable that fuscus was based on a dark
coloured specimen and represents no more than an individual
variation.
“ Native name, ‘Sikwikle.’
‘Very common and frequenting the native lands and the
adjacent bush.”—C. H. B. G.
26. Mus ratrus L.
6. 1520, 1526, 1527, 1551, 1581, 1652, 1657, 1658. 9. 1528.
‘¢ Native name, ‘ Tikonso.’
‘¢ Very common and confined to the kraals and houses, never
apparently visiting even the adjacent cultivated land. Several
other examples besides the specimens sent were seen and all were
very smallin size. Apparently exclusively nocturnal.”—C. H.B.G.
27. Mus coucHa Smith.
6. 1497, 1502, 1511, 1512, 1605, 1624, 1666. 9. 1480,
1504, 1648.
‘¢ Native name, ‘Supwisne.’
‘¢Common and habits similar to those of members of the species
in other parts of 8. Africa.”—C, H. B. G.
28. Mus cHRYSOPHILUS de Wint.
ge Lo0d, Lol. tol3) 1529, 15485 L60l) Os 1530) 1540)
Toa AG a a0:
‘Native name, ‘ Sinse.’
““Common ; habits similar to those of members of the species
in other parts of 8. Afvica.”—C. H. B. G.
29. TTHAMNOMYS ARBORARIUS Peters.
6. 1604, 1643. ©. 1644.
‘¢ Native name, ‘ Shikoloveta
“« According to the natives this species is fairly plentiful, although
they were unable to get me specimens ; the three secured being the
only examples I saw. It is arboreal in habits, ferming, according
to native report, nests similar to Dendromus or Th. dolichurus.”
—C. H. B.G.
30. LEGGADA MINUTOIDES A. Sim.
3. 1490.
“‘ Native name, ‘ Senbendenyumbe "
‘“‘ Apparently rare, as the specimen sent was the only one taken
or observed.”—C. H. B. G.
31. CRICETOMYS GAMBIANUS ADVENTOR, subsp. n.
6. 1632, 1641, 1651. 2. 1642, 1650, 1659.
Size and general characters as in gambianus and g. viator; the
296 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 19,
body-colour above much as in viator. ‘The sharp line of demareca-
tion between the colour of the back and the white of the belly,
which is strongly pronounced in typical gambianus, and plainly
though less markedly so in viator, completely absent; the cheeks
and flanks lighter than in viator, the lower part of the former white
like the belly. Otherwise the colour-pattern quite as in viator.
Skull in size and other essential characters as in viator ; but the
upper molar series rather stronger, the anteorbital foramina
slightly broader, and the rudimentary postorbital processes more
distinct.
Dimensions of the type :—
Head and body 358 mm.; tail 438; hind foot 71; ear 40.
Skull—greatest length 75 mm.; basilar length 65; zygomatic
breadth 36; nasals 3211; interorbital breadth 11°8; palatal
foramina 8°6; length of upper molar series 11.
Type. Adult male. B.M. no. 6.11.8.125. Original number
1632. Collected 14 Aug. 1906.
These specimens are from the 8.E. limit of the range of the
species, and it is consequently not surprising to find such small
differences as those recorded above between this and the Nyasa
form, from which it is separated by nearly two degrees of latitude.
‘“‘ Native name, ‘Sigwinye.’
‘“‘ Fairly common and generally found in thickets and densely
wooded places, where it forms burrows of 2-6 holes, almost
indistinguishable from those of Pedetes cafer. It is exclusively
nocturnal in habits, and usually is only to be obtained during
the dark phase of the moon; on moonlight nights it seldom comes
out, owing, according to the natives, to its great fear of the larger
owls. During the dark nights it lays in a store of food to tide it
over the moonlight nights, until it can again venture out. I have
observed that it also loosely fills the entrance of the burrow with
dead leaves during the time the moon is visible. Itis a vegetarian,
and in search of food often climbs shrubs and small trees. A great
article of food with the natives.”—C. H. B. G.
32. PEDETES GAFER Pall.
3. 1569 GQmmature).
‘“‘ Native name, ‘ Masengwi.’
‘‘ Not very common, and apparently somewhat locally distri-
buted. Habits similar to its congeners in other parts of 8. Africa,
forming the usual burrows in the more open forests, never in
the thickets where Cricetomys gambianus is usually observed.
They were extremely wary and difficult to trap and bad luck was
experienced with them, inasmuch as in several instances only the
toes were left in the trap and in one case the black end of the
tail. ‘ Majengwi’ was the name given to this species by the natives
at the Klein Letaba, hence this isan added proof of the existence
of the Springhaas in that loeality.”—C. H. B. G.
1907. MAMMALS FROM COGUNO, INHAMBANE. 297
)
33. LEPUS CAPENSIS AQUILO, subsp. n.
3. 1518, 1542, 1563, 1565. ©. 1568, 1667, 1669.
A Hare of the capensis group with the nuchal patch grey, and
the chin, chest, and belly snowy white.
Fur long, soft, and very fine, length on middle of back about
25 mm. General colour of upper surface “ broccoli-brown ”
tinged with ‘“drab-grey,” becoming lighter and gradually suffused
with butty on the flanks. Individual hairs divided into five rings
of the following colours and approximate dimensions :—proximal
ring “grey no. 9,” 7 mm. in length; second ring very pale ecru-
dvab, 5mm.; third ring jetty black, 4 mm.; fourth ring light
pinkish buff, 6 mm.; distal ring black, 3mm. Individual hairs
of belly about 30 mm. in length, soft, silky, and entirely snow-
white. Nostrils and lips edged with white, muzzle and vibrissate
area between wood-brown and isabella colour; a broad line ex-
tending from upper edge of orbit to within half an inch of muzzle,
and thence spreading downwards over lower region of cheek sandy
ervey; infraorbital area between wood-brown and isabella colour.
Eyes ringed with white; forehead coloured like back. Ears dis-
tinctly smaller than in the other subspecies, with the proectote*
coloured like forehead and back, fringed with yellowish-buff hairs
intermixed with white; metentote sparsely covered with minute
white hairs, outer fringe snowy white, absent terminally ;
metectote white, the terminal margin black. Nape-patch between
“ smoke-grey”” and “drab-grey.” Interramia white. Throat-
patch “pinkish buff.” Chest and belly snowy white. Outer side
of fore and hind limbs deep buff, becoming lighter and interspersed
with white on fore and hind feet. Tail short, compactly haired,
not loose and straggling as is generally the case in specimens of
ochropus, black above, white below.
Skull as in true capensis.
Dimensions of the type (measured in the flesh) :—
Head and body 472 mm.; tail 95; hind foot 113; ear 92.
Skull—greatest length 85°5 mm. ; basilar length 70; zygomatic
breadth 40-1; nasals, oblique length 38:2; brain-case breadth
28°0; diastema 24-4; palatal length 36-0; palatal foramina 22-7 ;
upper molar series 14:7; antero-posterior diameter of bulla 11:3.
Type. Male. B.M. ne. 6.11.8.132. Original number 1565.
Collected 6 July, 1906.
This subspecies may be distinguished from Z. ochropus by its
white chin and chest and the buffy colour of its flanks, in contrast
with the yellow of the latter on the sides and nape. It can be
easily separated from Lepus capensis centralis and L. c. granti by
their having a vinaceous buffy or pinkish buffy chest and flanks.
It bears more superficial resemblance to ZL. c. capensis however,
but is distinguishable by having a white chest and chin and a
white border surrounding the tip of the ear, instead of, as is usual,
* Thomas, P. Z. S. 1905, vol. ii. p. 359 (1906).
298 ON MAMMALS FROM COGUNO, INHAMBANE. [ Mar. 19,
the terminal portion edged with black. A comparison with a
considerable series of all the forms mentioned above shows that
examples of the present subspecies have a considerably shorter
tail and ears than is usual in any of them. Further material will
probably show that intergradation takes place between this form
and Z. ochropus, in which case the latter will take its place in
the group as a subspecies.
‘“¢ Native name, ‘ Nfundla.’
‘‘Gommon everywhere, especially in the valley of the Inyasuni.
Generally lying up during the daytime in the clumps of small
bush or grass, and feeding throughout the night, when they can
be taken with a noose on the numerous footpaths. Numbers are
caught by the natives in this way, the animal being with them a
staple article of food.”—C. H. B. G.
34, PoraAMOCH@RUS CHGEROPOTAMUS NYAS Maj.
Q. 1870.
Tn its strong ferruginous colouring this specimen quite agrees
with examples from Zomba, B. C. Africa, to which Dr. Forsyth
Major has given (P. Z.8. 1897, p. 367) the subspecific name nyase.
“Native name, ‘ Ngulubi.’
‘Plentiful in the thickets and dense forests, but extremely
difficult to secure. Always observed in pairs, and more or less
nocturnal in habits. _When pursued, they savagely attack the
dogs, repeatedly charging until killed by the hunters.”—C. H. B.G.
35, CEPHALOPHUS NATALENSIS A. Sm.
3. 1613, @. 1485, 1675.
‘“¢ Native name, ‘ Mungulwi’ or ‘ Munguli.’
‘‘Not common, and found in the dense forest and thickets,
which the ‘ Nhlengane’ loves; even there it is locally distributed,
seeming to confine itself to certain patches, from which it never
wanders far. Only one or two others, besides the specimen sent,
were observed, they were not however secured.”—C. H. B. G,
36. CEPHALOPHUS GRIMMI L,
@. 1503, 1582, 1640.
‘“‘ Native name, ‘ Munti.’
‘“¢ Not too common, and found both in the river-valleys and the
forests, visiting the Kaffir lands at night. Numbers of this and
other buck are taken by the natives for food.”—C, H. B. G.
37. RAPHICERUS NEUMANNI CAPRICORNIS Thos. & Schw.
Ss WL, USGA, Sy Ibeyet)
These specimens quite agree with those from Klein Letaba,
N.E. Transvaal.
‘““ Native name, ‘ Isipenu.’
“ Fairly plentiful but locally distributed, confining itself to the
more open forest and plain, along the river-valleys, away from
habitations. Generally observed in pairs.”—C. H. B. G,.
nS Ore) MR. R. I. POCOCK ON PALLAS’S CAT. 299
38. NESOTRAGUS ZULUENSIS Thos,
@. 162 Wore) 9% 15255 1576; 16225
The examination of these specimens makes it clear that the
subspecies zulwensis Thos. of the Nyasan livingstonianus is quite
constant in the essential characters in which it differs from the
latter, and we think ourselves justified in regarding zulwensis as a
distinct species.
‘“‘ Native name, ‘ Nhlengane.’
“Very common at Coguno and the whole country inland, but
unknown in the immediate neighbourhood of Inhambane. It
inhabits the dense thickets and undergrowth, where it has regular
tracks, and is generally observed in twos, and occasionally threes.
It can be easily obtained in the early mornings and late afternoons,
when it is found out feeding in the more open patches of bush
and along the Kaffir footpaths that intersect the thickets in every
direction, or by joining a hunt which the natives organise for the
special purpose of taking this species for food. It is an extremely
difficult buck to see when standing in the thickets, and can often
be heard giving a goat-like snort of alarm, although quite invisible
at only a few yards distance. The does greatly outnumber the
bucks, and it is impossible to make out the sexes in their habitat ;
a great proportion of the former are killed, and very old examples
of the latter are not often obtained.”—C. H. B. G.
39. CERVICAPRA ARUNDINUM Bodd.
Go MSGI WEBB Sy GEO.
“‘ Native name, ‘ Mhlangu,’
“Observed only in the river-valleys, in many of which they
are very plentiful, often being seen six and eight together. In
the valley of the Inyasuni, where they are little disturbed, they
feed throughout the day, and are remarkably tame. Individual
males vary much in the thickness of the neck.”—C. H. B. G.
April 9; 1907:
Dr. Heyry Woopwarp, F.R.S., Vice-President, in the Chair. «
Mr. RK. I. Pocock, the Superintendent of the Gardens, exhibited
a photograph and the skull of a specimen of the Manul or Pallas’s
Cat (Felis manul) that had recently died in the Society’s Menagerie,
and made some remarks on the species.
The specimen (text-fig. 87) was received in exchange from the
Zoological Gardens in Calcutta, in April 1906, and died from
broncho-pneumonia in April 1907. It was alleged to have come
from Tibet. Untortunately no exact locality was recorded; but
since the coloration of the skin and the structure of the skull
agree with those of examples from Tibet that have been described,
300 MR. R. I. POCOCK ON PALLAS’S CAT. [ Apr. 9,
there is no reason to doubt that the animal was captured in that
country *.
When alive, this Cat differed markedly, both in behaviour
and appearance, from most captive examples of the genus /elis.
The latter, if they happen to be tame, usually evince grati-
fication of notice by rubbing, with tail erect, against the bars
of the cage. When wild, they are either contemptuously in-
different to friendly overtures or receive them, crouched in a
corner, snarling. The Manul, on the contrary, although not tame
enough to be handled or touched, nor sufficiently friendly to rub
himself against the bars, showed no fear of spectators and no
wish to avoid them. He would boldly and aggressively but
silently advance to the front of the cage and, standing on his hind
legs, grip the bars with his fore paws, ready to scratch at a
confiding hand unwarily placed within reach.
Text-fig. 87.
Felis manul.
(From a photograph of a specimen living in the Society's Gardens.)
Like all the small Cats, he was usually very silent. He was
never heard to utter the growling snarl and guttural expiratory
hiss with open mouth so familiarly associated with irritation
* Except that the orbits are incomplete behind, the skull closely resembles that of
a speciinen from Ladak in the British Museum.
1
1907. | MR. R. I. POCOCK ON PALLAS’S CAT. 301
of temper im Cats; and the “spit” was a short, sharp sound
like “ts,” “ts,” “ts,” projected through nearly closed lips.
According to the keeper, Dixon, the “mew” or “ caterwaul ”
was a sound somewhat recalling a combination of the bark of a
small dog and the “hoot” of an owl. This was heard on one
occasion in reply to the typical “ caterwaul” of a female Uganda
Cat (/. ocreata), when “on heat.”
The tail was almost invariably carried with its posterior half
upcurled, so that the broad black confluent stripes on the under
side of its distal end were in full view from behind. The
conspicuousness of this jet-black area was enhanced in a marked
degree by his peculiar habit of jerking the end of the tail smartly
up and down. The lowness and width of the summit of the head
and the lateral “set” of the ears imparted to this Cat an aspect
totally different from that of all other species (text-fig. 87). In the
latter the inner edge of the ear normally rises obliquely outwards
from the top of the head to form, with the outer border, a con-
tinuous curve or anacute angle. Butin /’. manul the inner border
lies normally in the same plane as the top of the head, and meets
the vertical outer border at a right angle. This border rises from
the head at a point on a level with the outer canthus of the eye.
Since depression of the ears in Cats is an infallible sign of anger
or of predatory excitement, the simulation of this act caused by
the low and lateral setting of these organs in the Manul imparts
to his face a permanent look of ferocity and unrest, quite unlike
the placid aspect of other Cats with their ears normally erect.
So far as the Manul is concerned, one practical result of the
lowness of the forehead and the lateral setting of the small ears
is the power to peer over the edge of an object, like a rock or a
fallen tree-trunk, without depressing and closing the ears, and
without showing so much of the cranium as most of the “ high-
headed ” Cats do when so occupied. Other Cats, when watching
prey from behind some such point of vantage, always lower the
ears so as to make them invisible and, at the same time, partially
close them in such a way that quickness of hearing must be
interfered with to a greater or less extent. In this, perhaps,
may be found the explanation of the peculiar structural features
in the head which give the Manul its remarkable physiognomy.
Be this as it may, the above-proffered explanation was forcibly
suggested by the observation of the living Manul peering over
the edge of his sleeping-box and showing a relatively small
enon of head above the eyes, the ears at the time being
scarcely perceptibly depressed and not in any sense closed.
It has been stated by Gray *, on the authority of Hodgson,
that the pupils of the eyes are linear and erect. This was not
the case in the Society’s specimen. Under the influence of
sunlight the pupil contracted to a small circular or subcircular disk.
The ivis was yellowish.
* P. Z.S. 1867, p. 275. This statement probably misled Elliot into having the
eyes of F. manul in his monograph drawn like those of a domestic Cat.
Proc, Zoou. Soc.—1907, No. X XI. 21
302 MR. R. I, POCOCK ON PALLAS’S CAT. [ Apr. 9,
There was no marked seasonal change in colour, the coat
merely becoming thicker in winter than in summer.
Satunin has recently made Felis manul the type of a new
genus, Zrichelurus (Ann. Mus. St. Pétersb. ix. pp. 495-506,
1905), being the first to point out in detail the structural
peculiarities of the species. Unfortunately he overlooked the
fact that Severtzow had already proposed the name Otocolobus for
the same species (Rev. Mag. Zool. x. p. 386, 1858).
To Satunin also belongs the credit of showing that three
distinct. forms of Manul are recognisable, each typical of a par-
ticular geographical area. To two of these he gives subspecific
and to one specific rank.
Substituting Otocolobus for Trichelurus, his classification is as
follows :—
la. Otocolobus manul Pall. (typical form), from Trans-
caspia, Turkestan, and Siberia to the west of Lake
Baikal.
b. Otocolobus manul mongolicus Sat., from Mongolia and
Siberia to the east of Lake Baikal.
2. Otocolobus nigripectus Hodgs., Tibet.
I have not seen skins of the typical or of the Mongolian forms,
but, judging from Satunin’s description and the published figures,
I should say that the differences between them and the Tibetan
form are only of subspecific value. The name for the latter
therefore will be Otocolobus manul nigripectus.
The Society’s specimen of this subspecies presented the following
characters :—
Prevailing colour of face grey: some buff below and at the
sides of the nose: the eyes surrounded by a greyish-white area
which is bordered bya black streak above and below and partially
on the inner side, giving a characteristic spectacled look to the
face. The buff area round the nose set off by a black patch,
whence arise some black moustachial bristles; the rest of these
bristles mostly white and arising from blackish lines on the
whitish area of the upper lips. Some black spots running into
abbreviated lines on the cheek below the eyes. The two genal
stripes broad and jet-black, descending obliquely downwards and
backwards, the inferior arising from a spot on a level with the
middle of the eye, the superior from a point near the outer canthus
of the eye; area above the latter stripe grey, below the former
pale grey turning to white posteriorly ; the area between them
pale greyish white. Posteriorly the two stripes are confluent,
and merge below the ear with the sooty-brown hue of the throat
and chest. The dark hue of these areas relieved by the long
white tips to the hairs. Summit of head black speckled with
white; its fore part or the area in front of the ears marked with
some small asymmetrically disposed jet-black spots interspersed
with a few greyish-white spots. Back of ears greyish, passing
1907. | MR. R. I. POCOCK ON PALLAS’S CAT, 303
into black towards the head and with a darker rim; hairs on the
inner side and in front of the ears whitish grey. Prevailing hue
of the dorsal area of the neck and body silvery- or iron-grey, the
hairs sooty black basally, then white witha black tip*. Laterally
the dark-tinted basal portion of the hairs is paler in hue and less
in extent, the greater portion of the hair below the white sub-
apical band being whitish and strongly or faintly tinged with buff,
the buff tint showing up strongly when the hairs are parted. On
the lower side of the body the hairs have no black tint and the
buff tint disappears, leaving the belly and chest white but for a
clouding of blackish blotches on the chest behind and between the
fore legs. On the lumbar and sacral regions of the back there are
traces, mostly very faint, of narrow transverse black stripes.
The largest and most distinct of these lies about midway between
the shoulder and the root of the tail; on the right side it
measures about 57 mm., on the left about 50, the two being
separated by a median area of about 20mm. ‘This is the only
stripe that is evident on the right side. On the left side there is
one short stripe in front of it, and two abbreviated stripes behind
it, one low down on the side, the other higher up and just
traceable to the middle line. Behind this there are, on the
middle line of the back, three faint transverse blackish blotches,
probably representing the dorsal end of stripes. The tail is
greyer and paler than the back ; in its distal half there are three
well-marked black stripes, narrow dorsally and laterally, but
expanding and forming a triangular patch inferiorly where they
meet. The last stripe fuses laterally and inferiorly with the
terminal black tip; and since the expanded areas of the other
stripes are only separated by the narrowest intervals, it follows
that the distal half of the tail is practically black below. The
proximal half of the tail is marked above with three narrow
indistinct stripes, which, however, widen and become much more
strongly defined below. Fore leg whitish grey turning to creamy
buff on the paws; the brachial stripe distinct: other stripes
represented by blackish patches. Hind legs with thighs grey
like the body, and indistinctly spotted ; front of the leg greyish,
turning to pale buff on the paws; back of the leg up to the
hock rusty cream, a black patch on each side of the paw above
the median pad.
Measurement of dressed and flat skin :—Head and body 20 inches
(=500 mm.), tail 9 inches (= 225 mm.).
It is noticeable that the hair of the ventral surface, both on
the throat, chest, belly, and thighs, is considerably longer than on
the dorsal and lateral surfaces of the body. On the spinal region
it measures about 23 mm., and on the belly about 43 mm. The
coating of long hairs below is perhaps an adaptation to life in
* Satunin (op. cit. p. 497) says that the hairs on the back and sides have white tips.
Did he overlook the slender black tip ?
21*
304 MR. R. I. POCOCK ON PALLAS’S CAT. | Apr. 9,
Text-fig 88.
WAN
\)
\\
Skull of Felis manul, viewed from above. Nat. size.
cold latitudes, where lying or sleeping in the snow is possibly not
an uncommon occurrence, In that case the hairs would act as a
protection against chill to the delicate internal organs, especially
the intestinal portion of the alimentary canal *.
The skull of 7. manul has been described briefly by Milne-
Edwards and Blanfordt, and more in detail by Satunin, who gives
measurements of three examples. The skull of the specimen that
lived in the Gardens (text-figs. 88, 89) does not apparently differ
greatly from these. The chief peculiarities of the skull may be
shown by comparing it with skulls of Felis sylvestris and Felis
ocreata, since £, manwl is, im my opinion, an aberrant form of
the group exemplified by these two species.
* The ventral development of the hair in the Yak (Bos grunniens), also a denizen
of cold countries, is a parallel case. The size of the tutt of hair at the end of the
tail inthis animal perhaps acts as a protection against frost-bite of a part of an organ
where the circulation is weakest.
+ Rech. Mamm. p. 226. ¢ Mamm. Brit. India, p. 83.
1907. | MR. R. I. POCOCK ON PALLAS’S CAT.
Text-fig. 89.
Skull of Felis manul, side view.
Width across zygomata .........
behind postorbital processes ...
Ot: DLAI CARL ae ene up arabian
lessee DELYVEEMPOLDITS ye eisen er:
», of mesopterygoid fossa
Length of palate ..........
29
PT ROLEZ UE OMe Crate en eeyenee ree
Height from coronoid to angular process
Length of upper carnassial
PP OL LO Wer CALNASsIAl ones enes
Median length of nasals .........
distal edge
just above constriction
Shope! Merayeisler Ose jSebUUl | iaedsanopngoon panos nondon
Jebyse yl Memmye*sioi Coe SVC a jenn en oogbed Indnad asuro anal
across postorbital processes ............
| ., ot muzzle at base of canines ............
» of muzzle at infraorbital foramina
| ,, between auditory orifices Ae
») ©ACKOSS Uppel Ccarnassials’..................
from palate to occipital foramen ....
» Of mandible from condyle...............
Width of nasals in line with transverse
across constriction .........
FE. manul EF. sylvestris
(Scotland).
(Tibet).
85
67 |
69 |
55
41
A7 |
19 |
23
Py |
23 |
41
12
31
36
A3
56
25
Nat.
AY
tT
size.
U2? Mh
EAH ay
305
96
82
69
49
34
47
20
25
28
34
39
12
35
45
48
64
30
105
Td
| FP. ocreata |
(Suakin).
99
83
72
54.
36
47
i belay
Nek
28
32
40
12°5
36
46
50 |
65
28
115
75
306 MR. R. I, POCOCK ON PALLAS’S CAT. [ Apr. 9,
Some of the principal differences not shown in the above-given
measurements may be described as follows :—
é. Upper carnassial without inner lobe; no maxillary excres-
cence bounding the infraorbital foramen above and out-
side; malar sending up a long narrow process in front of
and considerably above the lacrymal foramen; infero-
anterior edge of orbit circularly rounded ; upper edge of
orbit elevated, higher than median portion of frontal bone ;
facial portion of skull abruptly inclined; occipital crest
small; interparietal and parietal crests absent ; temporal
crests marked by two shallow grooves nearly evenly
converging from the postorbital processes to the inter-
parietal; the smooth median and rougher lateral (tem-
poral) area of the parietals slightly elevated and separated
by a shallow depression ; fronto-parietal suture strongly
angled behind the postorbital processes and procurved
(concave forwards) dorsally ; basisphenoid longitudinally
BMCHOG RA as feratetanr ou caciben be cia eRe eran eer aces manul.
b. Upper carnassial with strong inner lobe; a distinct excres-
cence on the maxilla bounding the infraorbital foramen
above and externally ; upper process of malar not extending
past the lacrymal foramen ; infero-anterior edge of orbit
more ovally rounded ; upper edge of orbit depressed, lower
than median portion of frontal bone; facial portion of
skull evenly curved downwards from the frontal to the
middle of the nasals; occipital and interparietal crests
well-developed ; temporal crests, when not mesially con-
fluent, represented by a low ridge defining a median lyrate
area forming a practically continuous curve with the
temporal portion of the parietal bone; fronto-parietal
suture forming a nearly straight transverse line; _basi-
sphenoid nearly flat, only lightly arched longitudinally.
sylvestris and ocreata.
The skulls of two Tibetan specimens that I have seen do not
agree with Milne-Edwards’s statement that the nasals in Mongolian
specimens are strongly compressed in their posterior and corre-
spondingly dilated in their anterior portion. Satunin also states
that the nasals in the Tibet Manul (/. nigripectus) differ from
those of the typical form in being constricted in the middle and
distally expanded. In the skull of the animal in the Society’s
Collection, the constriction is very slight, amounting to only | mm.
in transverse width across the two bones. From this constriction
the bones expand very gradually forwards and backwards. ‘They
may be described as being of the broad type, such as is shown in
F’, sylvestris.
The infraorbital foramen is small and vertically oval, its
greatest length being less than the distance between its upper
extremity and the superjacent edge of the orbit.
iw Ha
S
t
a]
“GOlIaS ae “1lOnhea sheave l
“duit uesan'p UAT TSP elseeg YW
TWAS Wel OG S)
MN
4
‘duit wsex) “Pp
UONOTSISL SLOMNIUSN SOURS A.
“UAT esp eTTeeS
let Ww
1907. | ON FISHES FROM THE EASTERN TRANSVAAL. 307
The following papers were read :—
1. On a small Collection of Fishes made in the Eastern
Watershed of the Transvaal by Capt. G. H. Bruce.
By G. A. Bounencsr, F.R.S., F.Z.8.
[ Received March 12, 1907. ]
(Plates XVIII. & XIX., and Text-figures 90, 91.)
When recently serving in the Transvaal, Capt. G. E. Bruce,
5th Mounted Infantry, was so kind as to respond to an appli-
cation I made to him in 1905, to preserve some of the fishes.
in which he was interested, and which I felt sure would prove
of considerable scientific value, so little being known of the
piscine fauna of the Transvaal. I have now the pleasure of
drawing up a list of the species represented in a small collection
made by Capt. Bruce and presented by him to the British
Museum.
The rugged nature and muddy water of the Transvaal rivers,
together with the steep banks overgrown by reeds and rushes,
make it very difficult to collect, except by fishing with fly or bait,
and the laws against netting and dynamite ase very strict.
Further, the size of the collecting-jars at Capt. Bruce’s disposal
precluded the preserving of large specimens. Notwithstanding
these restrictions, the collection contains examples of several
species not previously recorded from the Transvaal, and of five
which are here described as new.
CHARACINID&,
1. Hyprocyon LinzeAtus Blkr.
The ‘“ Tiger Fish” occurs in the Inkomati and Krokodil Rivers,
and appears to be plentiful in all the rivers of the East Coast.
from the Zambesi to Swaziland.
2. ALESTES IMBERI Peters.
New to the Transvaal, where it was found in the Inkomati
River at Komati Poort. First described from the Zambesi, it.
has since been found in Lake Nyasa, in German East Africa, in
the Congo, and in Angola.
3. MICRALESTES ACUTIDENS Peters.
Inkomati River at Komati Poort. Not previously known
from south of the Zambesi.
CYPRINID&,
4, BARBUS RAPAX Stdr.
Three specimens from Groot Olifant River agree well with
Steindachner’s description and figure of a specimen from the
308 MR. G. A. BOULENGER ON FISHES [Apr.9,
Limpopo River, except for the presence of two pairs of barbels.
I cannot help thinking that the absence of anterior barbels in
the type specimen is due to an anomaly, or that their presence
has been overlooked.
5. Barus EuTzNIA Bler.
Several specimens from Klein Olifant River. This species,
originally described from Angola (B. kessleri Gthr. nec Stdr.), has
recently been rediscovered in N.W. Rhodesia by Mr. Neave.
6. BARBUS POLYLEPIS, sp. n. (Text-fig. 90.)
Depth of body 41 times in total length, length of head 33 times.
Snout rounded-subacuminate, 3 times in length of head, feebly
projecting beyond mouth ; diameter of eye 41 times in length of
head, interorbital width 33 times; mouth small, inferior, its
width 4 times in length of head; lips well developed, lower
continuous across chin; two barbels on each side, subequal in
length, 3 diameter of eye. Dorsal III 8, last simple ray strong,
e
bony, not serrated, its rigid part # length of head; free edge of
Text-fig. 90.
——— S CO
CROOK
ORY
Barbus polylepis.
the fin strongly emarginate; its distance from posterior border
of eye equals its distance from caudal. Anal IIT 5, longest ray
3 length of head, not reaching root of caudal. Pectoral 3 length
of head, not reaching ventral; latter below anterior rays of
dorsal. Caudal deeply forked, with pointed lobes. Caudal
peduncle twice as long as deep. Scales 43 oh 5 between lateral
Ime and ventral, 18 round caudal peduncle. Olive-grey above,
white beneath.
Total length 120 millim.
A single specimen from Klein Olifant River.
In the Groot Olifant River, Capt. Bruce obtained a large Barbus,
weighing 32 lbs., and too large for him to preserve, which, from
the notes he has taken, probably represented the adult of the fish
here described. Capt. Bruce counted 42 scales in the lateral line,
84 in a transverse series above the lateral line, and 6 between
the latter and the ventral fin. This large specimen further
differed in the much longer rays of the anal fin.
1907. ] FROM THE EASTERN TRANSVAAL, 309
7. BARBUS HOLUBI Stdr.
Groot Olifant River.
This fish may be the same as B. natalensis Castelnau, in-
sufficently described.
8. BARBUS BRUCH, sp. n. (Plate XVIII. fig. 1.)
Depth of body 33 times in total length, length of head 4 times.
Snout rounded-subacuminate, 2? times in length of head, strongly
projecting beyond mouth ; diameter of eye 5 times in length of
head, interorbital width nearly 3 times; mouth small, inferior,
its width 43 times in length of head; lips strongly developed,
lower continuous and forming a rounded mental lobe; two
barbels on each side, anterior not quite 3 diameter of eye,
posterior 3. Dorsal IV 9, last simple ray strong, bony, not
serrated, its rigid part 3 length of head; free edge of the fin
strongly emarginate; its distance from centre of eye equals its
distance from caudal. Anal III 5, longest ray 2 length of head,
not reaching root of caudal. Pectoral ? length of head, not
reaching ventral; latter below middle of base of dorsal. Caudal
deeply forked, with poimted lobes. Caudal peduncle once and
:
% as long as broad. Scales 29 z, 2 between lateral line and
ventral, 12 round caudal peduncle. “ Muddy grey above, white
beneath; dorsal yellowish grey; caudal and paired fins bright
pink.”
Total length 150 millim.
A single specimen from the Groot Olifant River.
9. Barsus sEcTOR, sp.n. (Plate XVIII. fig. 2.)
Depth of body 33 times in total length, length of head 44 times.
Snout rounded, 3 times in length of head, strongly projecting
beyond mouth ; diameter of eye 4 times in length of head, inter-
orbital width 2? times; mouth inferior, feebly curved, its width
3 times in length of head, lower jaw with a sharp edge; lower
lip restricted to the sides; two barbels on each side, anterior
about 3, posterior about 4 diameter of eye.. Dorsal IV 9, last
simple ray strong, bony, not serrated, its rigid part 2 length of
head ; free edge of the fin feebly emarginate ; its distance from
centre of eye equals its distance from caudal. Anal III 5,
longest ray # length of head, not reaching root of caudal.
Pectoral a little shorter than head, not reaching ventral; latter
below middle of base of dorsal. Caudal deeply forked, with
pointed lobes. Caudal peduncle once and # as long as deep.
Scales 29 Z, 23 between lateral line and ventral, 12 round caudal
peduncle. “ Muddy grey above, white beneath ; dorsal yellowish
grey; caudal and paired fins bright pink.”
Total length 140 millim.
A single specimen from the Groot Olifant River.
But for the shape of the head and the structure of the mouth,
owing to which it should be referred to the genus Capoéta of
310 MR. G. A. BOULENGER ON FISHES [Apr. 9,
Giinther, this fish could not be distinguished from the preceding.
Similar cases of close resemblance between species thus referable
to Barbus and Capoéta have been pointed out by me when
describing collections from Morocco and Kast Africa.
10. BARBUS ELEPHANTIS, sp.n. (Text-fig. 91.)
Depth of body equal to length of head, 4 times in total length.
Snout rounded, 3 times in length of head, projecting beyond
mouth; diameter of eye 4 times in length of head, interorbital
width twice and 3; mouth inferior, feebly curved, its width
33 times in length of head, lower jaw with a sharp edge; lower
lip restricted to the sides; two barbels on each side, anterior
= diameter of eye, posterior as long as eye. Dorsal IV 8, last
simple ray strong, bony, not serrated, its rigid part 3 length of
Text-fig. 91.
Barbus elephantis.
head; free edge of the fin strongly emarginate; its distance
from posterior border of eye equals its distance from caudal.
Anal ITT 5, longest ray 2 length of head, not reaching root of
caudal. Pectoral ? length of head, not reaching ventral ; latter
below middle of base of dorsal. Caudal deeply forked, with
©
pointed lobes. Caudal peduncle once and % as long as deep.
Scales 37 2 3 between lateral line and ventral, 14 round caudal
peduncle. ‘‘ Muddy grey above, white beneath; dorsal yellowish
grey ; caudal and paired fins bright pink.”
Total length 155 millim.
A single specimen from Groot Olifant River.
Very similar to the preceding species, but distinguished by longer
barbels and smaller scales.
11, Barus trimacutatus Peters.
Groot and Klein Olifant Rivers and Inkomati River.
12. Barsus INERMIs Peters.
Klein Olifant River. .
Previously known from the Zambesi only.
13. VARICORHINUS BRUCI, sp. n. (Plate XIX.)
Body strongly compressed, its depth 33 times in total length ;
1907. ] FROM THE EASTERN TRANSVAAL. 311
length of head 42 times in total length. Snout rounded, broader
than long, 4 length of head; eye lateral, 43 times in length of
head, twice in interorbital width; no conical tubercles on the
head ; mouth feebly curved, its width 3 times in length of head ;
two barbels on each side, anterior 4 diameter of eye, posterior as
long as eye. Dorsal TV 9, last simple ray strong, bony, not
serrated, its rigid part 3 length of head; border of fin concave ;
longest ray a little shorter than head. Anal IIT 5, not reaching
root of caudal. Pectoral a little shorter than head, not reaching
ventral, which is inserted below middle of dorsal. Caudal
deeply forked, with pointed lobes. Caudal peduncle twice as long
as deep. Scales 31 a 24 between lateral line and ventral,
12 round caudal peduncle. “ Back muddy brown, belly white,
fins grey.”
Total length 170 millim.
A single specimen from Klein Olifant River. ‘“ Lives in deep
still pools or under big rocks.”
Closely allied to the recently described V. ansorgii Blgr., from
the Kwango River (Congo System) in Angola, which has a deeper
body, a shorter caudal peduncle, and the eyes turned more
upwards. This is the fifth known African species of Varicorhinus.
SILURIDA.
14. CLARIAS GARIEPINUS Burchell.
Groot Olifant River.
ANGUILLIDA.
15, ANGUILLA BENGALENSIS Ham.-Buch.
Groot Olifant River. The specimen is referable to 4. labiata
Peters.
CICHLIDA,
16. HApPLocHROMIS MOFFATS Casteln.
Groot Olifant River.
17. Ti~aPIA NATALENSIS M. Web.
Inkomati River, near Komati Poort.
18. TILAPIA SPARRMANI A. Smith.
Groot and Klein Olifant Rivers.
EXPLANATION OF THE PLATES.
Pratt XVIII.
Fig. 1. Barbus brucii, p. 309.
2. ar sector, p. 309.
a. Mouth seen from below.
PuatEe XIX.
Varicorhinus bructi, p. 310, with upper view of head and mouth
seen from below.
312 MR. T. A. COWARD ON THE [| Apr. 9
On the Winter Habits of the Greater Horseshoe, Rhino-
lophus jferrum-equinum (Schreber), and other Caye-
haunting Bats. By T. A. Cowarp, F.Z.8.
[Received February 7, 1907. ]
Jn my paper “On some Habits of the Lesser Horseshoe Bat,
thinolophus hipposiderus (Bechstein)” (1) I referred to several
points of interest in connection with that species during its
winter retreat, and suggested that its congener, the Greater
Horseshoe Bat, Rhinolophus ferrum-equinum (Schreber), would
be found to have similar habits. Since writing that paper I
have had the opportunity of observing the Gr eater Horseshoe in
winter in caves in the Mendips, and am fortunate in being able to
compare my notes with those made by Mr. Bruce F. Cummings,
of Barnstaple, Devon, who at the same time was making
observations on the species in the neighbourhood of that town.
The conclusions that I arrived at require some modification, but
only one of them was not fully confirmed: I said that the bats
“usually” occupy different retreats in summer and winter; it would
be more correct to say that at times and in certain localities different
retreats are used at different seasons, for in the Cheddar caves the
bats occupy apparently similar positions in summer and winter.
I have, indeed, little knowledge of the habits of the bats in this
locality during the summer months, but I found the remains of
insects which must have been captured in summer lying beneath
places which the bats occupy in winter.
My observations were made at Cheddar, Somerset, and in the
neighbourhood, between December 27th, 1906, and January 7th,
1907, and, later, on a few examples in captivity. Mr. Cummings
paid a number of visits to some disused manganese mines in the
Pickwell Down Sandstone near Barnstaple between December 24th,
1906, and January 23rd, 1907, and also kept one bat alive for some
days. At the end of December and beginning of January I
found Greater Horseshoes both in the deepest recesses of the
caves—some of which are of great length—and close to the
entrances, even within reach of daylight.
I regret that I have no accurate record of the temperature in
the open at Cheddar during my visit, but on the first few nights
there were sharp frosts; later the weather was mild and at
entrances of the caves the temperature varied from 40° to 43° F.
Within the caves—at some distance from the entrances—the
temperature was fairly constant; in different spots [ found it
between 50° and 52° F., and this I understand is also the summer
temperature. Mr, Cummings also found that the temperature in
the Barnstaple mines was 52° 10
The dimensions of the caves which I visited vary considerably.
The Long Hole, in which JI found the Greater Horseshoe
abundant, is a long natural tunnel in the limestone of perhaps 150
or 200 yards in length, and with an average width of seven or
1907. | WINTER-HABITS OF CAVE-HAUNTING BATS. 313
eight yards and a height of from five to ten feet; in places the
roof is much higher and culminates in fissures and hollows which
afford suitable hiding-places for large numbers of bats. The floor
is uneven and partially blocked by ancient roof-falls. At the end,
beyond a considerable mass of fallen rock, the cavern opens into
a small chamber, from which a chimney, blocked by a huge “‘ chock
stone,” runs upward to a chamber of considerable size. Out of
this upper chamber a low passage, in places less than eighteen
inches high, extends for some yards; the end of this passsage 1s
blocked, and when I examined it no flicker was noticeable in the
flame of my candle, but [ was informed that at one time it was
possible to feel a current of air near its termination, as if it led to
some outlet. From the lower chamber a lime-encrusted shaft,
running upwards, may lead to further chambers or to the open.
During a portion of my visit I was accompanied by Mr. Charles
Oldham.
In the Long Hole I found Greater Horseshoes occupying a
fissure high up in the roof near the entrance. I cannot say how
far within this fissure the bats rested in the daytime, for they
were not visible; it was only after dusk that I saw them emerge
from some hole in the fissure. I found other Greater Horseshoes
scattered singly in the main tunnel, hanging from the roof or in
the water-worn holes with which it is abundantly pitted. In
the terminal chamber the bats were in colonies of varying size.
In one of the caves of the village, which is open for visitors and
illuminated by gas, there is, near the entrance, a wide shaft open
to the air. Some ten feet from the floor of the cave, and perhaps
fifteen feet from the top of the shaft, a small side-chamber opens
out of the wall; the floor of this chamber was thickly covered with
old dry dung, mingled with fragments of many beetles, chiefly of
the genus Geotrupes. Dr. J. Harold Bailey, who kindly examined
all my coleopterous fragments, identified portions of Geotrupes
spiniger Marsh, and of another Geotrupes. The Greater Horse-
shoe, as I shall show later, captures some, and in summer probably
all, of its prey outside the caves and conveys it to the interior of
the caves before consuming it. The head, elytra, and other
portions of the beetles are dropped whilst the bat is crushing the
insect in its jaws. his is undoubtedly a common habit, but bats
which go far from the caves in search of food may hang to
convenient footholds at some distance from their diurnal sleeping-
places. In captivity the bats rest after a meal and frequently
defecate, a habit which probably accounts for the piles of dung
beneath favourite feeding-places in the caves. Both Mr. Cummings
and I noticed that our captive bats showed a marked preference
for certain resting-places, settling again and again within a few
inches of the same spot. This chamber in the shaft was perhaps
the only place I came across which appeared to be solely a summer
haunt; it may or may not be occupied as a sleeping-place in
summer, but it was undoubtedly a chamber to which numbers
retired to feed.
314 MR. T. A. COWARD ON THE [Apr. 9,
Great Oons Cave is longer than the Long Hole; it consists of
a rough, slightly descending tunnel and a steep passage running
downwards to a considerable depth from the end of the main tunnel.
At the bottom of this terminal passage, at a great distance below the
entrance of the cave, I found one Greater Horseshoe hanging ;
a few others and Lesser Horseshoes, Rhinolophus hipposiderus
(Bechstein), were in other parts of the cave, and a Whiskered
Bat, Myotis mystacinus (Leisler), was close to the entrance. The
Kcho Cave is a great cleft in the cliff, entered by a small hole and
short passage; the cleft then extends upwards and downwards, the
height being over a hundred feet and the width little more than
eight feet. Thirty feet or more above the floor of the cave
colonies of Greater Horseshoes were hanging from the wall on
one side, and a few Lesser Horseshoes were suspended, singly,
from the opposite wall. Goatchurch Cavern, on the northern
slope of the Mendips, has its entrance high on the hill-side in the
treeless valley of Burrington Combe. This cavern consists of long
and intricate passages, running far into the hill and descending
steeply; here again at a great depth L found both species of
Horseshoe scattered about the passages. In other smaller and
shallower caves, some with narrow entrances and others with wide
mouths, I found Greater and Lesser Horseshoes; one of the
former hung in full daylight ten yards from the entrance, and a
Lesser Horseshoe from the upper part of a bottle-shaped hollow
in the roof only four yards from the entrance.
In no ease in which I was able to make repeated observations
were the bats actually hibernating. All were susceptible to the
disturbance caused by my entrance, drawing themselves up by
bending their legs and often swaying slightly, although not
touched ; bats changed their positions in the intervals between my
visits, while others, as I shall show later, flew in the caves
without any artificial stimulus and indeed went out into the open
air of their own accord. Lesser Horseshoes and Whiskered Bats
also moved from positions in which I had observed them, and I
saw a Long-eared Bat, Plecotus auritus Geoftr., on the wing in
the evening in one cave.
On December 29th I carefully noted the position of two Greater
Horseshoes in the main tunnel of the Long Hole, one of which
was not more than ten yards from the entrance, but was careful
not to touch them nor to bring my light neartothem, Twodays
later both had moved. On the same date (Dec. 29th) I found a
colony of about forty hanging close together—only a few were
actually touching any of the others—in the terminal chamber.
Five of these I took without touching any of the others. When
pushed from their footholds they fluttered to the ground and lay
there, cold and lethargic, but squeaking feebly and extending their
legs backwards, apparently in search of some foothold. The
remainder drew themselves up by bending their legs but did not
unfold their wings. Very little dung lay beneath this colony, and
the bats which I took, though they were soon awake and lively,
1907. | WINTER HABITS OF CAVE-HAUNTING BATS. 315
did not defecate in the box; one taken near the entrance of the
cave did so at once.
On the 29th I went into the cave in the afternoon, but on the
31st, when I found that the two bats whose positions I had noted
had moved, I entered about 6 p.m. A few yards from the
entrance, in a fissure in the roof which may lead to chambers and
passages as yet unknown, a thick cluster of wide-awake Greater
Horseshoes was suspended from a spur of rock. On the floor of
the cave beneath this fissure I had, on the 29th, noted a large
pile of dung, but certainly, during the daytime, no bats were
visible in the fissure. The bats, which numbered at least thirty
and possibly as many as fifty, were clinging together like a swarm
of bees; their heads, ears, and facial ornaments were in constant
nervous movement and the wings hung loosely by their sides;
they had apparently emerged from some crack or hole in the
fissure. When I turned the light of a powerful lamp upon the
bunch, individuals at once detached themselves, dropped on
outstretched wings, and flew further into the cave. On this
occasion, so far as I could tell, all flew into the depths of the cave
and not towards its mouth. J struck two down with my stick as
they flew and could have killed many more; within afew minutes
the fissure was empty, ail having apparently entered the cave.
As I proceeded up the tunnel I found many bats clinging
to the walls and roof, their wings hanging loosely, their heads,
ears, and nose-ornaments twitching, as with raised heads and
necks and hollowed backs they turned and twisted in all
directions without moving the position of their feet. Whether
these wide-awake bats were or were not able to see me I am
not prepared to say, but they were keenly alive to my
presence and moved, flying freely, when I approached them.
Other bats passed me, flying up and down the tunnel, but it is
probable that there is either a second exit or further chambers,
for I certainly did not come across all the bats which had left the
cluster near the entrance. In the terminal chamber, where on
the 29th I had seen about forty bats, only eight remained, and
these again were awake; while I was watching them two took wing.
On January Ist, 1907, I went in in the daytime, and not only
found that the six bats left on the evening before in the terminal
chamber had disappeared but could not see a single bat hanging
in the cave. On January 3rd I entered about 6.15 P.m.; no bats
were visible in the fissure near the entrance where I had seen the
cluster on December 31st, nor were there any hanging and oniy one
was on the wing in the tunnel. The chamber at the foot of the
chimney was again empty, but by climbing over the chock stone
I reached the upper and larger chamber which I had not previously
visited. Here I found two small colonies or collections of bats—
numbering twelve and eight individuals respectively—and a
single bat in the low passsge which leads out of this chamber.
These bats were not in profound slumber; some two or three
individuals were hanging with the wings by their sides and were
316 MR. T. A. COWARD ON THE -Avor. 95
in nervous motion. I took the single bat, but did not touch nor
otherwise disturb the colonies.
On January 5th, with Mr. Oldham and my usual assistants I
entered the Long Hole about 5.20 p.m.; bats were then dropping
out of the fissure near the entrance singly, but there was no cluster
as there had been on December 31st. About half a dozen came out
and flew inwards. A few were on the wing in the tunnel and the
two colonies in the upper chamber were reduced to one company
of ten individuals. We again left this colony undisturbed.
In the afternoon, on January 6th, we again visited the upper
chamber and found that the number of the bats was further
reduced; only four remained. No bats were flying in the cave
between 3 and 4 P.M.
We then returned to the fissure, where I had seen the cluster
and whence on the previous days we had both seen bats emerge. At
4.20 p.m. a single Greater Horseshoe dropped out of the hole and
flew into the cave, and at 4.40 another dropped from the fissure
and flew out into the open air. Mr. Oldham then posted himself
at the entrance to the cave, in such a position that he could
command a view of any bats passing out, while I remaimed below
the fissure. Between 4.40 and 5.20 p.m. at least nine bats came
from the fissure or elsewhere and flew out of the cave. A few
bats passed us in the tunnel; we could hear the whirr of their
wings distinctly, and before it became too dark to distinguish
objects at a distance of a few feet—at about 5.30—bats passed.
and repassed the entrance to the cave, perhaps some of those which
had emerged intending to re-enter. So long as there was light it
was easy to see the broad wings of the Greater Horseshoes, as they
passed out or fluttered past the entrance, silhouetted against
the sky.
My visits to the other caves were not so frequently repeated.
On December 29th I did not observe any Greater Horseshoes in
Great Oons Cave, but I noted the position of two Lesser Horseshoes
and a Whiskered Bat without touching them ; the Whiskered Bat
was clinging to the roof within full reach of daylight. One of the
Lesser Horseshoes was sleeping lightly; it partially unfolded its
wings, raised its head, and moved its ears, and then relapsed into
slumber, refolding its wings. Not one of these three bats, on
January 5th, was occupying the position I had seen them in on
December 29th, but one Lesser Horseshoe and a Whiskered Bat
were in places which were not occupied on the previous visit.
We found three Greater Horseshoes, two of which were not at the
spots they occupied on January 5th, when I visited the cave on
December 29th.
One large cavern was visited on several occasions; on De-
cember 31st I could see a number of Greater Horseshoes hanging
singly in different parts of the cave, but I only took or touched
one; on the subsequent visits I did not see a single bat.
About 7 p.m. on December 31st there were at least thirty or
forty Greater Horseshoes, hanging in two companies some
1907. ] WINTER-HABITS OF CAVE-HAUNTING BATS, 317
distance apart, from the wall of the Echo Cave; the majority of
these bats were awake and others were on the wing. On
January 5th the number was smaller; most of these bats were so
close together that six were dislodged at once by a stone thrown
from below. The Lesser Horseshoes in this cave may or may not
have moved in the interval between the visits; they were roughly
in the same position on both dates. A Long-eared Bat, which,
when handled and replaced in the same position, appeared to be
in a deep sleep on the 31st, had gone by the second visit. An
unidentified bat was on the wing in the daytime just within
the entrance of Wookey Hole, a cave near Wells.
Between December 24th, 1906, and January 23rd, 1907,
Mr. Cummings paid seven visits to the old mines near Barnstaple ;
he arrived independently at the conclusion that the bats were
not hibernating and were going outside the caves to feed. He
has kindly allowed me to add his notes.
His observations were made at three borings. On December 24th
he found a single Greater Horseshoe in No. 1, but none in No. 2;
No. 3, the floor of which was under water, was not entered. On
December 29th No. 1 was unoccupied, but there were two bats in
No. 2 twenty-five yards from the entrance. The bats were not
touched, but when, later in the afternoon, he re-entered this
boring, he found one on the wing, and the other shortly followed
the first out of the tunnel. On January 5th there were again two
bats in No. 2. One occupied a similar position to that when first
noted, but the other had not returned to the same position. The
temperature in the open, on the date of this visit, was high, but
rain was falling steadily. The two bats, after Mr. Cummings had
taken them out of the tunnel to examine them, and had then
returned them to their original positions, were roused and driven
out of the cave; they attempted to re-enter the cave three times,
but were apparently prevented from doing so by Mr. Cummings’s
presence at the entrance. Mr. Cummings thought they objected
to the rain. Later in the afternoon he discovered two bats, which
showed by their manner of hanging with their wings by their
sides that they were not in deep sleep; he feels sure they were
the bats driven out of No. 2. On January 11th No. 1 and No. 2
were empty, but a small bat entered No. 3 at about 5.20 p.m. and
shortly afterwards two Greater Horseshoes came out; Mr. Cum-
mings had, again, not been able to enter No.3. On January 16th
Nos. | and 2 were again unoccupied ; the temperature in the open
was on this and the previous visit 48° F.; within the caves 52°
and 53° F. At 5.30 p.m. two Greater Horseshoes, which came
from the direction of a deep shaft higher up the slope, passed the
entrance of No. 38. On January 19th, when the temperature in
the open had fallen to 40° and a cold east wind was blowing, no
bats were seen at the mouth of No. 3 between 5.10 and 5.45 p.m.,
neither were any noticed between 5.15 and 5.45 p.m. on
January 23rd, when the temperature at the mouth of the cave was
33°, and it had been below freezing-point all day in the open.
Proc. Zoot. Soc.—1907, No. XXII, 22,
318 MR. T. A. COWARD ON THE [Apr. 9,
Mr. Cummings, on this evidence, suggests ‘That the Greater
Horseshoe, at the end of December and throughout January,
moves freely in the caves, is easily disturbed, and usually hangs at
the further end of the boring. That it goes abroad regularly in
mild weather, but hibernates more or less deeply in frost, and
remains in the caves during rain or high wind.”
It has been often asserted that in the Greater Horseshoe and
other gregarious bats the sexes hibernate apart; though colonies
consisting of bats of one sex only have been found occasionally,
both sexes may be found in the same company or cluster. I have
found male and female Lesser Horseshoes in the same colony at
Cefn, Denbigh, and of the five bats which I took from the forty
in the Long Hole on December 29th one was a female and four
males. The two bats which I killed as they left the cluster at the
mouth of this cave on December 31st were males; four taken from
the upper chamber of the Long Hole on January 6th, the remnant
of the company of twenty, were maies; six dislodged by a stone
from a congregation on the wall of the Echo Cave were all males;
and five Pipistrelles, Pipistrellus pipistrellus (Schreber), turned
out of a crack in the cliff-wall, outside the entrance to the Long
Hole, were also males. The single bats found in different places
were of either sex, but the majority were males; their ages
varied and there were many which, judging by their grey pelage,
were born in 1906. I may, of course, have failed to discover the
colonies of females, or some of the bats which we did not touch in
the colonies may have been of this sex; there is, however,
another possible explanation, an inequality in the numbers of the
sexes, the males predominating.
The Greater Horseshoe feeds in winter, apparently both within
and outside the caves. The floors of all the caves show evidence
of the recent movements of bats ; in addition to the piles of dung,
which were mostly old and dry, fresh excrement was scattered, as
if dropped by the bats either when flying or hanging from the
roof. Fragments of the prey, in places in small heaps, were also
littered about the floors of the caves.
The stomachs of two Greater Horseshoes, killed immediately
after capture, contained no food, but that of a third, killed as it
flew from the bunch at the entrance to the Long Hole, contained
a little and there was feecal matter in the intestine.
Large beetles and moths are apparently the chief food of the
Greater Horseshoe. I found elytra of MJelolontha vulgaris Fabr.,
and the heads and elytra of Geotrupes spiniger Marsh, and perhaps
G. stercorarius Linn. ; these remains were in most of the heaps of
dung and were also scattered in the caves; in the Great Oons I
found them more than 100 or 150 yards from the entrance. Both
species of Geotrupes are occasionally abroad in mild weather in
winter : we found one G. spiniger on horse-dung in the road near
the caves; Mr. Cummings finds Geotrwpes in small numbers
throughout the winter at Barnstaple. In the Cheddar caves were
the elytra of the flightless beetle, Vebria brevicollis Fabr., and at
1907.] WINTER-HABITS OF CAVE-HAUNTING BATS. 319
Barnstaple Mr. Cumimings found fragments of a flightless Ptero-
stichus (possibly, he thinks, P. niger Schall.). The remains of the
large cave-spider, Weta menardi Latr.—fragments of the cephalo-
thorax with the legs attached in some cases—were present in the
Cheddar caves, and Mr. Cummings found remains of a spider at
Barnstaple; at Cefn, I found a portion of the leg of a Meta in
the dung of the Lesser Horseshoe (1). These flightless beetles
and spiders must have been picked up by the bats from the ground
or taken from the walls. It is not certain that the bats always
secure their prey when they are flying, and as the Horseshoes are
exceedingly agile on the wing I feel sure that they could, by
hovering, pick food from the ground or from a vertical surface,
such as the wall of a cave, without alighting ; furthermore, in
captivity they show a marked tendency to fly low, and will fre-
quently alight with outspread wings upon a flat surface, springing
therefrom again with ease.
The wings of Scotosia dubitata Linn., a moth which hibernates
freely m limestone-caves and was abundant at Cheddar, were
scattered here and there, as if eaten recently, and also a few wings
of Gonoptera libatrix Linn., another species which hibernates,
were present. These moths might, of course, have been captured
when they were on the wing in summer, but the remains appeared
fresher than those of summer-flying moths which do not occupy
the caves in winter (for example, Zriphena) which I found.
Mr. Cummings found the remains of both these moths at Barn-
staple, and also fragments of two large flies, possibly Hristalis
tenax and some smaller dipteron. At Cheddar there were many
small dipterous insects resting on the walls of the caves; these are
soon aroused to activity by the presence of a light. Myr. Robert
Newstead found, im dung which he kindly examined for me,
remains of certain J/uscide, which, as he points out, are all diurnal
insects. We do not, however, know at what hours the bats are
on the wing in summer, but even if the times when the bats
emerge and the insects retire in the evening, or vice versd in the
morning, do not overlap, the bat which can capture a flightless
beetle or spider could take a fly which was at rest.
In Goatchurch Cavern, at a great distance from and below the
entrance, I found a dead Staphylinid beetle of the genus Quedius ;
it was damaged by the teeth of a bat.
The prey of the Greater Horseshoe may be captured on the
wing, but that it is not, as a rule, devoured whilst the bat is flying,
seems to be proved by the behaviour of bats in captivity even
more than by the presence of the fragments of prey in the caves.
When secured by a snap of the bat’s jaws the insect is conveyed
to some resting-place and there consumed. ‘Two captive Greater
Horseshoes, one of which survived for 14 days and the other
for 35 days, were fed almost entirely upon Greotrupes typheus
Linn., the only beetle I could obtain in sufficient numbers in
winter. Altogether over 120 beetles were devoured, and in every
case the behaviour of the bats was practically the same. I usually
20%
320 MR. T. A. COWARD ON THE [Apr. 9,
held the bat in my hand until it had snatched the beetle and
then released it; but at times the bat, when suspended from
some foothold, would take a beetle which was offered to it. The
released bat, holding the beetle securely, even if only by one leg,
invariably flew to some favourite foothold—usually the picture-rail
in one of three different spots, one end of the window-sill, or a
curtain-ring—and there hung until the beetle was devoured.
When the beetle was dropped in flight, only two or three times
in over 120 experiments, the bat made no attempt to recover its
prey. I never heard the sound of champing jaws as the bats were
flying, but when they were at rest the noise of crushing the hard
armour of the beetles was plainly audible.
The interfemoral membrane was never used as a pouch, as it 1s
in the Vespertilionide (2), but the beetle was invariably pushed
against the interbrachial membrane, as I observed was the case in
the Lesser Horseshoe (1). At times it was thrust against the
belly, sometimes into one and sometimes into the other wing, and
as a rule one leg was detached from its hold in order to give more
freedom to the half-outstretched wing on the same side. The
contortions of the bat were most noticeable when it was hanging
clear and not against the wall; on many occasions one or other of
my captives went through the whole operation when suspended
from my finger. The object of this use of the membrane is un-
doubtedly to prevent the escape of the struggling beetle as the
bat relaxes its grip in order to adjust the position of the captive
in its mouth ; when the beetle was seized by the head, especially
if it was a male armed with sharp thoracic spines, it was turned
until the bat had a grip upon the abdomen. As the head is
thrust into the interbrachial membrane, the wing and leg, on the
same side, are moved sharply forward, thus, by the resistance of
the membrane, enabling the bat to secure a firmer grasp of the
beetle, almost as if the wing was used actually to push the prey
further into the mouth. The object of pouching in the Vesper-
tilionide is to prevent the escape of a captive, when the bat is
flying, until a firm grip is secured (2), but mm the Horseshoes the
use of the interbrachial membrane appears to be different ; in the
Vespertilionide the head is rapidly withdrawn from the pouch and
the prey devoured openly ; in the Ahinolophide the wing is made
use of repeatedly, so long, in fact, as any large portion of the prey
projects beyond the jaws of the hat.
The head, prothorax, first pair of legs, and elytra of a beetle are
usually dropped by the bat, but occasionally the head or one
elytron are devoured ; those portions rejected by my captives were
similar to many of the fragments found in the caves. When a
bat had eaten two or three beetles its head drooped and it lapsed
into partial slumber, rousing itself spasmcdically at any sudden
noise. After a few minutes, however, it would be lively again,
swinging round on its legs and moving its head and ears at the
slightest sound.
Mr. J. G. Millais states (8) that this species “ does_not devour
1907. ] WINTER-HABITS OF CAVE-HAUNTING BATS. 321
all its food while in flight, but conveys some to its diurnal resting-
place”: I will go further; it probably conveys all its food to
some resting-place. Even when I gave my captives mealworms,
insects which are easily crushed by the powerful jaws of a Greater
Horseshoe, they were conveyed to some resting-place unless the
bat refused to leave my hand. Mr. Millais adds a tootnote : “* Lam
quite ignorant as to the manner in which these bats carry their
food to their retreats, since the interfemoral pouch seems to be
incapable of being bent forward.” In no bats, to my knowledge,
is the pouch used as a receptacle in which to carry food; the prey
is carried, in the most natural manner, in the mouth, firmly secured
by the strong teeth. Mr. Millais’s statement (p. 30) that the tail
is bent forward and the interfemoral membrane occasionally used
is probably based on some erroneous observation ; it is the inter-
brachial and not the interfemoral membrane which is used. During
the whole process of pouching and eating the beetle the posterior
portion of the tail remains in the recurved position which is so
characteristic of the species.
This reflexed tail, or, to be more exact, portion of the tail, is
constantly in this curious position. In flight the anterior portion
of the interfemoral membrane is stretched between the slightly
flexed legs; the end of the tail is upturned; and when the bat
is scrambling or climbing the tail is held in the same position ;
when at rest the tail is flat upon the back if the wings are half
open, or lies partially concealed by the forearms if the animal is
closely wrapped in its wings.
In sleep the position of the wing-membranes is practically the
same as in the Lesser Horseshoe (4), but, asa rule, the bats I found
were not so completely encircled by the wings as is the case with
the smaller species; the forearms seldom met along the back,
whereas in the Lesser Horseshoe one sometimes overlaps the other.
In some of the sleeping Greater Horseshoes the tail stood out
from the back at an angle of about 30°, in a position I have never
observed in the smaller species. Our knowledge of the habits of
the Rhinolophide is slight, and as yet we cannot find that the bats
gain any advantage from this recurved tail ; there is, however, the
possibility that the tail is rudimentary.
Occasionally, im captivity, as Mr. Millais noticed, the Greater
Horseshoe will sleep with the wings folded along the sides, but
this is an unnatural position and is probably accounted for by the
impaired health of the captive; when in good health the bats
sleep with the interdigital membranes one above the other, in a
similar position to that adopted by some, at any rate, of the
Pteropodide. The ears, slightly bent or folded, are hidden beneath
the carpal joint.
The colour of the adult is brown and does not vary in the sexes,
but the young are grey; all the young which | found were
naturally much darker than the August young depicted by
Mr. Thorburn (5), having more nearly acquired the brown pelage
of maturity.
322 MR. T. A. COWARD ON THE [Apr. 9,
We find, to sum up, the following facts :—
1. The Greater Horseshoe Bat, and apparently in the mild
climate of Somerset the Lesser Horseshoe also, if the weather be
open at the end of December and beginning of January, is not in
a state of hibernation. It moves in the caves, awakening without,
artificial stimulus, and leaves the caves apparently in search of
food.
2. To a great extent the same haunts are used in summer and
winter, seeing that numbers of both species are to be found
hanging near the entrances to the caves in the winter in situa-
tions certainly resorted to during the summer.
3. Both species are susceptible to the presence of a man when
they are apparently asleep. They show by their movements—by
bending their legs and raising and slightly swaying their bodies—
that they are influenced by the disturbance. It appears, from the
way in which colonies dwindled in size from time to time, that
the bats, after being thus disturbed, though not actually awakening
at the time of the visit, retire from the situations they occupied
to deeper recesses.
4. Food is conveyed into the caves from without and devoured
there, the bats hanging whilst they feed.
5. Certain creatures are captured and eaten in the caves.
6. Creatures incapable of flight are captured by the bats and
devoured.
7. When feeding the Greater Horseshoe, like its congener,
makes use of the interbrachial membrane and not of the inter-
femoral pouch.
The following is the result of Mr. Robert Newstead’s examination
of the dung and insect remains which I obtamed in the Cheddar
caves.
Wings of the following moths :—
Scotosia dubitata Linn. | Probably captured when these insects
Gonoptera libatriz Linn. } — were hibernating in the caves.
Triphena orbona Fabr.
Triphena pronuba Linn.
NXylophasia polyodon Linn.
Captured and conveyed into the
caves in summer.
Dry dung, obtained in the summer haunt where I found no
bats in winter. 41 pellets minutely examined.
About 68 per cent. contained the remains of Lepidoptera.
ee OO - “ us Coleoptera.
PES ae es y : Diptera.
5 aaah: 3 s Arachnida (Spiders).
4 igs) 5 by Hymenoptera.
* Pagal tos “ A Trichoptera.
Of the Coleopterous remains at least 44 per cent. were of
Geotrupes ; in about 12 per cent. there were fragments of dMelo-
1907. ] WINTER-HABITS OF CAVE-HAUNTING BATS. 323:
lontha; in one pellet there were portions of the elytra of a
Geodephagous beetle, possibly of the genus Amara, and in another
what appeared to be Dytiscus.
The fragments of Lepidoptera were difficult to identify, but a
number of abdominal segments, scales, hairs, antennz, eyes, and
eges were from moths of the family Noetwidee.
One pellet was almost entirely composed of the remains of
Diptera. Mr. Newstead remarks, “including portions of the
wings of a large Muscid with metallic green body, possibly Lucilia
or allied genus.” ‘This green fly was pr resent in seven pellets; the
remains of other Diptera were in three or four pellets.
In one pellet was ‘“‘a portion of the wing of a Hymenopteron
remarkably like Vespa.”
One pellet was entirely composed of the remains of a caddis-fly.
In December and January I found a caddis-fly on the wing at
Cheddar, and I picked up the remains of one amongst the
fragments in the Long Hole.
The spider-remains were of a large species, probably the cave
Meta; 1 found remains of this spider, certainly dropped by the
bats, in the caves.
Mammalian haivs, probably those of a bat, were present in one
pellet.
In 100 pellets, taken from below the fissure in the Long Hole,
whence we watched bats emerge, about 48 per cent. of the remains
were of Geotrupes, and a smaller percentage of a large brown
species, possibly Melolontha. This dung was dry; we may
conclude, not only from the presence of a summer flying beetle
(presuming that it was Jelolontha) but from the condition of the
dung and the large size of the heap, that the fissure is occupied
in summer.
Fifty pellets taken from a shallow wide-mouthed cave and
evidently dropped recently, contained about 98 per cent. of metallic
fragments of the abdomen and basal segments of the legs of
Geotrupes.
One hundred recent pellets from the Long Hole were composed
of about 90 per cent. of remains of G'eotrupes.
It would thus appear that during the winter months beetles of
the genus Geotrupes form the chief food of the Greater Horseshoe
Bat, and that in summer Lepidoptera and Coleoptera are devoured
in, approximately, equal quantities.
The following parasites were on Bats which I obtained at
Cheddar :—
Nycteribia hermanni Leach. From two examples of Ahinolophus
hipposiderus. Identified by Mr. Percy H. Grimshaw.
Eschatocephalus vespertilionis (C. L. Koch). On &hinolophus
Serrum-equinum, Rhinolophus hipposiderus, and Myotis
mystacinus. The Ticks, all females, were identified by
Mr. A. 8S. Hirst.
324 MR. F. E, BEDDARD ON THE ANATOMY pavprend,
Spinturnix vespertilionis (Dufour). This small Mite was present
on the wing-membrane of Rhinolophus ferrwm-equinum, and
was identified by Professor E. Trouessart, of Paris, to whom
Mr. Hirst had kindly forwarded my specimens.
REFERENCES.
(1) Cowarp, P. Z.8. 1906, pp. 849-855.
(2) OupHAM, Zoologist, ser. 4, vol. ui. pp. 471-474 (1899).
(3) Minnats, Mammals of Great Britain and Ireland, vol. i.
Ds Ale
: (4) OtpHam, Manchester Memoirs, vol. xlix. no. 9, pp. 6-7
(1905).
(5) Miuuats, op. cit. vol. i. plate 1.
3. Notes upon the Anatomy of a Species of Frog of the
Genus Megalophrys, with references to other Genera of
Batrachia. By Frank EH. Brepparp, M.A., F.RS.,
Prosector to the Society.
[Received March 19, 1907. }
(Text-figures 92-100.)
I possess, through the kindness of Dr. Charles Hose of Borneo,
an example of a Frog from that country, which at first appears
to be referable to the genus Megalophrys, and which at least
comes nearest to—if it be not identical with—Ceratophrys
nasuta of Schlegel. It appears to differ, however, from that
Species in one very 1mportant point and in one or two features
of minor importance. The single example was in an excellent
state of preservation, and I am therefore able to contribute some
facts to the anatomy of this genus, which, save for many external
characters and certain osteological features, is apparently quite
unknown to Zoologists. It is necessary, however, to preface my
account of its anatomy with an attempt to fix accurately the
species to which this account refers. The species I. nasuta was
originally named and figured by Schlegel*. Our subsequent
knowledge of the external characters of the species is chiefly due
to Cantor7, Duméril & Bibron +, Giinther §, Boulenger ||, who
has summed up and added to the previous descriptions, and, latest
of all, Werner]. Darwin** has figured the heads of Megalophrys
* Handi. Dierk., Atlas, pl. iv. fig. 72. The leaf-like appendage upon the nose is
represented as bemg directed forwards, which is not the case with my example.
+ Catalogue of Reptiles inhabiting the Malayan Peninsula &c. Calcutta, 1847,
p. 140.
{£ Erpétologie Générale, vol. viii. p. 456.
§ The Reptiles of British India, Ray Soc. 1864, p. 413; Ann. Mag. Nat. Hist.
(4) xi. p. 419.
|| B. M. Cat. Batrach. Sal. 1882, p. 443.
{| “Reptilien und Batrachier aus Sumatra,” Zool. Jahrb. (Abth. f. Syst.) xiii.
p. 498. ** Descent of Man, 1871, i. p. 26, fig. 32.
1907.] _ OF A FROG OF THE GENUS MEGALOPHRYS. 325
nasuta and M. montana, regarding them, in accordance with the
first and erroneous view of Giinther, as male and female respectively
of Megalophrys montana. Kdeling* has figured the entire animal
from above. All of these descriptions, with the exception of
that of Boulenger, leave a good deal to be desired in point of
details of importance, doubtless because those details were
hardly appreciated as important at the time the papers or works
in question were written. To Mr. Boulenger’s definition, Werner
has added a more accurate account of the dermal ossifications,
the mere occurrence of which was known to his predecessors. I
shall have occasion to refer again to all the above-named writers
in the following statement of certain differences which the Frog
studied by me shows as compared with J/. nasuta.
The length of the Frog which forms the subject of the present
communication is 135 mm. from snout to vent, a measurement
which nearly agrees with that of Werner t, whose description is
not sufficiently comprehensive to allow of a confirmation of his
identification, The most salient external characters which point
to my Frog being identical with Megalophrys nasuta are, in
correlation with other characters, the long palpebral horn-lke
appendages and the leaf-like appendage upon the snout. The
tympanum moreover is concealed, and the loreal region 1s ex-
cavated. Furthermore, the conical warts agree exactly in their
disposition with the descriptions given by Cantor, Giinther, and
Werner ; that is to say, there are three, one on the sacrum median,
and a pair anteriorly in the shoulder-region. There are, more-
over, three warts upon the head, all of different sizes, arranged
in a triangle, the largest occupying the apex of the triangle
which is directed backwards. Ey (eling ¢, however, whose figure
of the “species” Megalophrys chysii is not very good, represents only
the sacral wart and nothing of the kind upon the head. Those,
indeed, seem to have also escaped the attention of others. No
characters of value are introduced into his two descriptions of
the species, and it is impossible to be certain of the identity of
“ Vegalophrys chysti.” She appendages over the eye and on the
nose are not of course diagnostic of the genus or species; for
in various Hemiphractide “there are similar appendages, and
this family is not to be confused with the Pelobatide. Moreover,
My. Boulenger has recently § removed from the genus Megalophrys
and assioned to the allied genus Leptobrachium, “Me galophrys fee ||,
a species with prominent palpebral processes, the existence of
which doubtless originally influenced Mr. Boulenger in placing
the species in that genus. It has also the hard skin upon the
back which is mentioned as being more strongly developed in
* Nat. Tijdschr. Nederl. Indié, vol. xxvii. 1864, p. 265 & plate.
+ Giinther also (Ann. Mag. Nat. Hist. ser. 4, xi. p. 419) speaks of the frog as
5 inches long. Flower, on the other hand (2: ZS. 1899, p. 916), gives 90 mm. as
extreme length.
t Ned. Tijdschr. y. de Dierk. vol. ii. 1865, p. 205, & Nat. Tijdschr. Nederl. Indié,
vol. xxvii. 1864, p. 265 with plate.
§ Ann. Mus. Genova, vol. vil. p. 750. || Zbe vol. v. p. 512.
326 MR. F. E. BEDDARD ON THE ANATOMY [Apr. 9,
M. nasuta than in M. montana. In the example studied by
myself, the calcified (?) area formed a continuous plate beginning
immediately behind the head, but not continuous with the
hardened skin lying over the head, reaching back rather beyond
the anterior ends of the ilia. Laterally this area is bounded
by a raised glandular fold. On the ventral side of this fold the
skin is still hardened, though not so much as dorsally ; and this
area is again bounded by a narrow raised area quite similar to
that just referred to, which separates it from the soft but tuber-
cular skin of the belly. The inner finger is distinctly longer
than the second, a character in which this species agrees with
M. longipes* rather than IM. montana‘.
It is part of the generic definition of J/egalophrys according
to Mr. Boulenger that the outer metatarsals are united. In
Text-fig. 92.
Palmar surface of hand (upper figure) and foot (lower figure) of
Megalophrys nasuta.
this it differs, according to the same author, from e. g. Pelobates,
where the “ outer metatarsals are separated by web.’ Having
been able to compare my Frog with Pelobates fuscus, | find that
* Boulenger, P. Z.S. 1885, p. 850. + Id., Cat. Baty. Sal. p. 442.
1907. ] OF A FROG OF THE GENUS MEGALOPHRYS. 327
both frogs agree in the separation by web, only that the web
is more extended towards the tips of the toes in Pelobates*. In
this, my species apparently agrees with Megalophrys longipes.
It has, however, a fairly conspicuous inner metatarsal tubercle,
but nowhere projecting from the surface of the foot as in
Pelobates. I found no trace of an outer metatarsal tubercle or
of subarticular tubercles. These are absent in JZ. longipes, but
described (together with the inner metatarsal tubercle) as “ in-
distinct” in MW. montana and (by inference) M/. nasuta.
There are other characters used for systematic purposes in
which the individual which I have dissected does not agree with
its presumed congeners. ‘The vomerine teeth, which are very
few 7 and form only a narrow band over the projecting region
of each vomer, are situated distinctly to the inside of and not
behind each choana. And the loreal region is deeply coneave, as it
is stated to be in Megalophrys montana and (by inference) in
M. nasuta by Boulenger =, but apparently not in MW, longipes.
As a part of the generic definition of M/egalophrys, Boulenger uses
the form of the tongue, which is described as “ subcircular,
indistinctly nicked and free behind.’ In my example of
Megalophrys the tongue is very faimtly$ nicked behind. It is
also nearly circular and free behind.
Very important also in fixing the systematic position of this
Frog are two osteological characters which have been largely
used in defining the genera of Batrachia. The fusion of the
sacral vertebra with the ensuing coccyx is a rare feature in
Batrachian osteology. It occurs, however, in Pipa and among
the Pelobatide. In two genera only, viz., Scaphiopus and Pelobates,
does Boulenger describe the sacrum and coccyx as confluent.
They are most unquestionably so in the Frog which I describe
in the present communication, and this confluence is incidentally
shown in the drawing (cf. text-fig. 93) on p. 332, illustrating the
arrangement of certain muscles. This fusion of the sacral
vertebra with the coccyx is not only an important classificatory
fact if existing systems are to be respected, but raises another
fact of greater Importance. It is well known that among those
genera of Anura in which this fusion occurs, viz. Pipa, Xenopus,
and Hymenochirus—the entire set of genera constituting the
Aglossa—and in Pelobates, Scaphiopus, and the present genus
among the Pelobatidee, and in Bombinator among the Discoglosside,
the apparently single sacral vertebra is really double in the case
of Pipa and Pelobates, formed of three vertebrae in Hymenochirus,
and of only a single vertebra in Scaphiopus and Bombinator. It
becomes, therefore, a matter of great interest to ascertain what
is the arrangement that characterises the sacrum of the species
* Giinther, Cat. Batr. Sal. 1858, p. 136, speaks of the toes of Ceratophryne nasuta
as “completely tree.”
+ Giinther, Cat. Batr. Sal. 1858, p. 1386, writes—“ Vemerine teeth none.”
{ Cat. Baty. Sal. p. 542.
§ “ Tongue entire behind.”—Giinther.
328 MR. F. BE. BEDDARD ON THE ANATOMY [Apr. 9,
which I provi isionally at least identify with Megalophrys nasuta.
The most obvious test to apply for the solution of this question
is the exit of the spinal nerves. Now I find that a stout nerve
issues from the spinal cord just in front of the stout and expanded
transverse process of the sacral vertebra; and that an equally
stout nerve issues behind this transverse process in the angle
between it and the coccyx. There is none between the two,
Hence the sacrum is composed of but one vertebra as in most
other Frogs.
A second character is the proccelous excavation of the vertebral
centra, A part of the definition of MJegalophrys by Boulenger is
“ Vertebree opisthocelian.” This character, used by him in the
tabular * discrimination of the genera. of ‘Pelobatidee, is again
made use of in the fuller deser iption of the genus Megalophrys *.
It is, however, in the characters of the sternum that this
species chiefly difters from other species of Jegalophrys. Among
the characters used by Boulenger to define the family Pelobatide,
one is stated as follows, viz.: “The omosternum is constantly
present, but small and cartilaginous.” That this definition is
not absolutely true was subsequently shown by Mr. Boulenger
himself §, who asserted that in Scaphiopus solitarius there was
“no omosternum.” The shoulder-girdle and sternum of the
genus Megalophrys have been figured and deseribed by Parker ||
in Megalophr ys montana. The omosternum is minute and the
sternum is ossified nearly throughout, an inconspicuous cartila-
ginous xiphisternum only being left at the free extremity of the
bone. These characters (¢nter alia) are used by Mr. Boulenger
in his definition of the genus Megalophrys, and therefore
presumably also apply to WW. nasuta. Whether the sternal
apparatus of JZ. longipes has been examined I do not know. In
the Frog upon which I report in the present communication, the
characters of the sternal apparatus are different from those
which have been referred to in MJegalophrys montana. Both
sternum and omosternum are very well developed.
The omosternum, instead of being a small oval plate of cartilage
as it is represented to be by Parker in We galophrys montana, has
form which apart from details is more like that of Rana
esculenta. It is actually ten millimetres long, and its proportions
to the rest of the sternum are therefore much more like those
of Paludicola bibronti figured by the same author . Although,
as in WW. montana and other Frogs, the procoracoids are bent very
* Cat. Batr. Sal. 1882, p. 433.
+ Ibid. Pp. 442 ; t Loe. cit. p. 432.
§ P.Z.S. 1899. p. 790.
|| A Monograph on the “Structure and Development of the Shoulder-Girdle,”
Ray Soc. 1868, p. 78, pl. vii. fig. 8.
{| It is by no means certain in every case that the species described by Prof.
W. K. Parker in his Monograph of the Shoulder-Girdle have been correctly
identified. I find for example that the sternum of Hyla caerulea (= Calamites
eyanea” of Parker’s nomenclature) is not as Parker has figured it (Joc. c?é. pl. vii.
fig. 6), but exactly resembles the sternum of “ Acrodytes daudinii” (presumably
a Hyla) figured by the same author (Zoe. cit. pl. vil. fig. 1).
€
1997. | OF A FROG OF THE GENUS MEGALOPHRYS. 329
much forwards, their articulation with the omosternum in JZ. nasuta
is not by any means so far forwards as it is figured by Parker
in WM. montana. his articulation is plain enough in M/. nasuta.
Above this articulation and for the entire shaft the omosternum
is clearly calcified. It expands above into a circular cartilaginous
plate which has a crescentic outline anteriorly and is wider than
the shaft. The procoracoidal cartilages at and near both points
of articulation with the omosternum are also calcified. The
sternum consists of a bony style as in Megalophrys montana; but
there are differences in detail. In the first place, the ene 1s
longer in proportion to its width in W. naswta, and it does not
expand so markedly in width towards its posterior termination
as is represented in Prof. Parker’s figure of Megalophrys montana.
In the second place, J/. montana is characterised by a cartilaginous
xiphisternum which “is but little extended either laterally or
axially beyond the shaft-bone.” This is not at all the case with
Megalophrys nasuta, where the xiphisternum is a broad and
expanded plate, having posteriorly the semicircular cheese-cutter-
like outline which is so usual among Frogs. On the whole,
therefore, there are some differences between the two species.
The sternum of this Frog is in fact particularly large as
compared, for example, to that of its ally Pelobates fuscus, with
which I have compared it in this and in some other details of
structure. The difference of size is, of course, actual in view of
the much larger dimensions of the Frog which I describe here as
compared with the rather small Pelobates fuscus. In Pelobates
the total length of the body in the individual measured was
47 mm. and the total length of the sternum 16 mm., the sternum
being therefore roughly one-third of the length of the body.
The corresponding measurements in the Frog described here
weve 135 mm. and 60 mm., the proportions therefore nearly
approaching one-half. The relationship of the sternum to the
underlying viscera shows corresponding differences in the two
genera. In Pelobates the sternum hardly extends back beyond
the heart and pericardium which, however, it fully covers. The
liver is left largely exposed. In ‘“ Megalophrys nasuta” the
sternum passes a considerable distance beyond the liver-lobes,
the heart being beneath almost the commencement of the sternum
proper.
So far as I can gather from the memoirs already quoted which
deal with the species Megalophrys nasuta, there has been no
actual description of the sternum in this particular species. But
since several systematists use the occurrence of a rudimentary
omosternum as a generic definition, the matter must have been
looked into by them, or by some of them. So far as present
views upon the classification of the Anura go, it is clear that I
should be hardly wrong in instituting not merely a new species
but a new genus for this Fr rog, on account of its diver gent sternal
characters,as compared with those that have already been described
in the genus Megalophrys. In contradiction, however, to this
330 MR. F. E. BEDDARD ON THE ANATOMY [ Apr. 9°
conclusion is the extraordinary similarity in detail-—I particularly
recall the three warty tubercles upon the back and the ‘“nose-
leaf”’—between this species and J/. nasuta; but coupled with
these are also, as it would appear, difterences in features of similar
ov nearly similar value. Thus the species described here differs
from J. nasuta, in that the vomerine teeth are within a line
joining the choanee, and that the inner finger is distinctly longer
than the second. Again, however, I infer these differences,
being unable to find any definite description of the characters in
Megalophrys nasuta.
The above statement concerning those characters used in the
discrimination of genera and species among the Anura seems to
me (unless very grave errors have crept into existing descriptions)
to show that quite possibly two species have been confused under
the name of JJegalophrys nasuta, which may even be legitimately
veferred to different genera. Ido not propose, however, for the
present to give a nametothe species the anatomy of which I deal with
in the following pages, in case systematists have really had before
them, and described, a Frog which is identical with it. But even
in this case the foregoing description of external characters is not
without use; for some of these characters have been undoubtedly
overlooked, or imperfectly described, as in the case of the sternum.
The above account of the systematic characters of the Frog
which, on a superficial examination, would be referred to Mega-
lophrys nasuta as described by Cantor, Gunther, Schlegel, and
Werner, and, I presume, Boulenger*, may be conveniently sum-
marised for future reference. The following is a restatement of
the characters of Megalophrys nasuta based upon the single female
example of a Frog from Borneo, which, from a survey of the
external characters as described by several zoologists, would be
referred to that species ;—
Length about 5 inches. Head broad and depressed, with loreal
region excavated. Tympanum invisible. Long palpebral process
over each eye and an upwardly directed leaf-shaped process on
the snout. Vomerine teeth few, between choane. Tongue sub-
circular, very faintly nicked, free behind. Teeth only on upper
jaw. Three inequisized warts forming a triangle upon posterior
region of head. Skin of back induraced, like that of head, toa
joint behind front ends of iia. This region separated by a narrow
elandular fold from lateral region, also, but more slightly,
indurated, which is again separated by a similar fold from the
warty ventral surface. Three prominent tubercles upon back, one
* Myr. Boulenger’s recent description (Ann. Mus. Genov. ser. 2, iv. p. 512) of a
Frog, referred to the genus Megalophrys, and to a new species of the same, viz.,
M. fee, which was later (ibid. vil. p. 750) removed by him to the genus Lepto-
hrachium (with which he also fused the genus Xenophrys), seems to show that
Mr. Boulenger had not before him at the time when he wrote his ‘ Catalogue of the
Batrachia Salientia’ examples of the Frog described by myself in the present
communication. For he has mentioned in describing “ Megalophrys” fee the
fact that the vertebree are proccelous and that they are opisthoceelous in Meyalophrys
nasuta.
1907. | OF A FROG OF THE GENUS MEGALOPHRYS. 331
on each shoulder and one on sacral region. Vertebre proccelous ;
sacral vertebra fused with coccyx, its transverse processes greatly
expanded. Noribs. Omosternum not rudimentary, with calcified
style; sternum a bony style, with expanded cartilaginous xipki-
sternum ; shoulder-girdle arciferous. Fingers free; inner finger
longer than second. ‘Toes half-webbed ; outer toe separated from
next by web. Toes not dilated at tips; no articular tubercles ;
inner metatarsal tubercle extensive but not separated anteriorly
from the surface of the foot ; no outer metatarsal tubercle.
Hab. Borneo.
It will be observed that in the above brief diagnosis I have not
attempted to differentiate between generic and specific characters
or indeed family characters. That this Frog belongs to the
Pelobatidee is quite plain. That it cannot be referred to any
known genus—if the present definitions of the same are retained—
is alsoobvious*. I propose, however, for the present to defer the
question of generic distinctness until the Pelobatide are better
known anatomically 7.
Whether this Frog is a new species or identical with Megalophrys
nasuta | must, for reasons already stated, leave uncertain.
§ Muscles of the Back.
In the various drawings (text-figs. 93-96) submitted herewith in
illustration of the diaphragm of different species of Batrachia, the
principal muscles of the back are also shown, and it will be at
once seen that these differ considerably in the different forms.
Only in Megalophrys is it impossible to see these different muscles
(text-fig. 98), for the lengthy transverse muscle of the cesophagus
and lung almost completely covers them over. When, however,
this muscle is divided along its greater length, 7. e. across the
direction of its fibres and the two flaps reflected, the muscles now in
question are fully displayed as shown in text-fig. 93. The same
muscles are present which are known to exist in Rana esculenta,
with the possible addition of a muscle not found either in
h. esculenta, R. guppy (see p. 333), or 2. tigrina, of which species
I have examined the last two. The Jntertransversarii muscles
are well developed and commence from the anterior margin
of the greatly expanded transverse process of the sacral vertebre.
They are so well developed that they almost conceal the anteriorly
lying transverse processes of the vertebrze, each separate band of
muscle joining successive transverse processes being attached near
to the ventral median line of the same. The Jlio-lumbaris muscle
* (Footnote added May 8th.) After the reading of this paper, Mr. Boulenger
kindly drew my attention to the fact that there occurs in Pelobates cultripes
a variation In connexion with the fusion or non-fusion of the sacral vertebra
with the coceyx (‘The Tailless Batrachia of Europe,’ Ray Soc. 1897, part i. p. 208).
It does not, however, follow that if a character is variable in one species it is not
a “ good ” character in another.
+ Schlegel’s name Ceratophryne cannot be resuscitated ; for he first applied it to
C. dorsata, an American frog which is, I presume, Ceratophrys dorsata.
332 MR. F. E, BEDDARD ON THE ANATOMY (Agr. 9%
is comparatively unimportant, when those muscles in some other
Frogs to be shortly described are considered. As shown in text-
fig. 93 its branches to the transverse processes are limited to the
outer edges of these. The enormously expanded sacral vertebree
which overlie the iha give off anter lorly the strongest slip of
muscle which Irefer to the Jlio- lumbaris * complex. <A slender
slip of muscle represents the origin of the muscle from the ilium.
This arises from the tip of the ilium as it has been described to
arise in Rana esculenta; but instead of running forward as in
Part of dorsal musculature of Megalophrys nasuta.
es. (Esophageal muscle cut and retlected with oviduct. Coce.I/. Ilio-coccygeal.
Ii.lumb. Long slip of Mio-lumbaris muscle compared in the text with the
Musculus proprius pulmonum of Pipa. 8S. Transverse process of sacral
vertebra, the fusion of which with the coccyx is shown. To the right of
the long ilio-lumbar slip three short ilic-lumbars are shown overlying the
Intertransversarii.
that frog, it runs dorsally at right angles to the longitudinal axis
of the body owing to the overlap of the sacral vertebra to which
it is partly attached. Jegalophrys differs, as will be seen presently,
from a number of other Frogs in the presence of a long, almost
strap-shaped and strong muscle which may perhaps be referred to
the /lio-lumbaris complex. This arises from the ilium a long way
back and is continued straight forwards, without any insertions
of detached slips on the way, to the third vertebra, to the outer
** This corresponds throughout the following descriptions to the “ Pars lateralis ”
of Gaupp.
1907. | OF A FROG OF THE GENUS MEGALOPHRYS. 333
cartilaginous extremity of the transverse process of which it is
attached. A flat slip of muscle connects the third vertebra with
the second, and is exactly in the same straight line with the
muscle just described. It gives the impression “of being a portion
of it although it is not actually connected, There is nothing
exactly com parable to this muscle in any of the genera of Batrachia
with which I shall presently deal, with the exception of the allied
Pelobates. Iam inclined, however, to think that it may be the
homologue of the “musculus pulmonum proprius” of Pipa*,
which the enormous growth of the transversalis in Megalophrys
has cut off from communication with the aponeurosis of the lung
and diverted to the transverse processes of adjacent vertebree.
But this is at present no more than a suggestion.
Text-fig. 94.
Some of the dorsal muscles of Rana guppyi.
T. Lung. o.d. Oviduct. V. Vertebral centra. Other lettering as in text-tig. 93.
A comparison between the muscles of Jegalophrys which have
just been described and those of Rana shows a great number of
differences. In Megaiophrys the Ilho-lumbar muscle complex is
* Beddard, P. Z.S. 1895, p. 839, fig. lim. The transversalis sheet to the lung is
not shown in this figure, ‘ret is in that of Keith (J. Anat. Phys. yxxix. p. 261,
fig. 13 c).
Proc. Zoou. Soc.—1907, No. XXITT. 23
334 MR. F, E, BEDDARD ON THE ANATOMY [ Apr. 9,
not important as compared with the Intertransversarii, that is if
the long muscle running from the ilium directly to the third
vertebra be put out of consideration. In Rana guppyion the other
hand (see text-fig. 94) the ilio-lumbar muscles are very important
and occupy a considerable space upon the transverse processes of
the ribs. The whole muscle appears to te continuous from the top
of the ilium to the transverse process of the vertebra. But it is,
though strictly continuous, interrupted by tendinous intersections,
each one of these corresponding to a transverse process. These
tendinous intersections are of some width. In Rana tigrina, which
I have been able to compare, the muscle has the same general
arrangement; but the tendinous intersections are hardly
noticeable. This difference is very possibly merely one of those
differences which may be fairly put down to size, In the larger
Rana guppyi the complication of the muscle is greater than in its
smaller ally.
Text-fig. 95.
ra
"e
GQ See il
A dissection of Pelobates fuscus, to show large esophageal sheet of the
transversalis. Lettering as in text-figs. 98, 94.
Pelobaies shows a distinct likeness to Megalophrys in the dis-
position of the muscles now under consideration, as might be
1907.) OF A FROG OF THE GENUS MEGALOPHRYS. 335
expected from other points of likeness between the two genera,
which are referred by some systematists to the same family of
Aveifera, The striking feature about the muscles of this region
of the body in Pelobates, is their very massive development as
compared more particularly with Bufo and Ceratophrys, which I
shall deal with later on in this communication. When the viscera
are pushed to one side to render visible these various muscles, the
ilium and the transverse processes of the vertebree are nearly
completely invisible. The latter indeed are entirely so; but a
portion of the ilium is exposed where the massive ilio-coccygeal
muscle diverges from it. This is by no means the case with other
genera, as the series of figures (text- figs. 93-96) illustrating these
facts shows. In this feature again Megalophrys comes nearer to
Pelobates. But in AMegalophrys the expanded sacral vertebre are
left uncovered; they are quite covered in Pelodates.
The strong J/o-coceygeal muscle covers nearly the whole of the
ilium, arising up to its very tip. To the imside, ¢. e. towards the
centre of the vertebral column, it conceals the commencement of
the intertransversarius. The latter muscle is also very strong
and thick. Instead of lying between adjacent transverse pro-
cesses, It is not on their plane at all, but entirely covers them and
moreover shows no visible tendinous inscriptions, Quite anteriorly
at the second or third vertebra, this muscle finally meets the
equivalent of the peculiar muscle described above in Jegalophrys.
That muscle, as in AMegalophrys, avises in Pelobates a long way
down the ium; it is strap-shaped, but relatively stouter than in
Megalophrys. In front of the iltum it had no attachment to—
sent down no slips to—the transverse processes of any vertebre
save the anterior vertebra to which it is ultimately attached. I
could detect no ilio-lumbaris other than this muscle, even on
removing it from its close contact with the intertransversarius ;
nor indeed is there much room for one.
So far as the material upon which Tam able to report here enables
me to say, Ceratophrys stands at one end of a series commencing
with Pelobetes, and of which Rane is an intermediate member in
so far as concerns the muscles of the back which are here dealt
with. Im Ceratephrys the muscular development of this region
of the body is but feeble, relatively speaking. The transverse
processes of the vertebree and the larger part of the ilium are
quite uncovered by muscles, which are extremely shrunken as
compared with those of Megalophrys and Pelobates and even Rana.
The same muscles precisely can be recognised and with but little
modification. The Iho-coceygeal is much reduced and arises only
from the internal edge of the ilium, commencing some way below
the tip of that bone and not extending on to the ventral side.
The Intertransversarii are quite separated by the successive trans-
verse processes. There is no continuous band of musculature as
in Pelobates and, though to a less extent, in some of the other types
described here. The Hio-lumbaris is peculiar as compared with
that of other genera: it arises as a thick band of muscle from
23%
336 MR. F. E. BEDDARD ON THE ANATOMY [Apr. 9,
near to the tip of the ilium. It forms a continuous mass without
tendinous intersections, giving off, however, slips to three trans-
verse processes only, upon the most anterior of which the muscle
ends.
Text-fig. 96.
Some of the dorsal muscles of Ceratophrys ornata.
Lettering as in text-figs. 93 & 95.
In Ceratophrys the vertebre thus suppled by branches of the
ilio-lumbaris muscles are Nos. 5, 6, 7; but a few fibres are seen
to run on to the transverse process of the fourth vertebra.
Bufo closely resembles Ceratophrys in all these features. The
ilia and the transverse processes of the vertebree are but little
covered with muscle—at any rate in Lufo marinus and B. agua,
the two species studied by myself. The ilio-coccygeal is, in pre-
cisely the same way, a much reduced muscle, commencing its
origin some way back upon the ilium and not extending upon the
ventral surface of that bone. The ilio-lumbaris is if anything a
rather slighter muscle than that of Ceratophrys. Otherwise it is
exactly similar in its appearance and relations to Bufo martinus
1907. | OF A FROG OF THE GENUS MEGALOPHRYS. 337
(I am not quite certain as to B. agua); its fibres are inserted on to
the transverse processes of vertebree 4 to 7 inclusive. It is from
the fourth vertebra in these species that the esophageal and lung
muscles arise (see below, p. 346). In ana also it is from the
fourth vertebra.
The resemblance between Bufo and Ceratophrys in the muscles
now under consideration, particularly in their feeble development
as compared with the corresponding musclesin some other genera,
might be put down to similarity of habit. For Ceratophrys 1s
emphatically a ‘toad’ in its terrestrial habits and lethargic life.
1 find, however, that there is a close likeness in respect of these
various structures between Ceratophrys and Leptodactylus which
is important in view of their inclusion in the same family Cysti
enathide by some systematists. JT have examined two individuals
of Leptodactylus hevadactylus, which agree entirely in all the
structural features to be referred to immediately. A considerable
portion of each ilium is left perfectly free of muscle. As in
Ceratophrys, but not in Bufo, the anterior extremities of the ila
do not reach as far as the transverse process of the eighth vertebra ;
they extend beyondthem in Lufo. The ilio-coccygeal commences
some little way down the ilium and gives off slips to the transverse
processes of vertebre 4 to 8 inclusive, and the origin of the lung-
muscle is confined to the fourth vertebra.
There is not, in fact, much difference in these particulars
between Leptodactylus and Rana. For though in the large Rana
guppy? the ilium is well covered by the ilio-coccygeal muscle, this
is not the case with Rana tigrina. And the well-marked tendi-
nous intersections of the muscle in the larger Rana are not seen
in the smaller species, which thus resembles Leptodactylus. In
Ceratophrys the modifications as to these muscles have gone still
further, and it is perfectly clear from the figures (text-figs. 94, 96)
that that genus shows marked, though in reality superficial
differences from other Frogs, which may perhaps, in view of its
likeness to Bufo, be looked upon as adaptive. It 1s, however,
perfectly clear that Megalophrys and Pelobates, though differing
from each other in various details, are on the whole much more
different from the other genera treated of here than are any of
those genera among themseives.
S$ Muscles of the Ventral Surface.
In the sternal region of MJegalophrys nasuta the following
muscles are displayed when the skin is removed :—(1) Pars
abdominalis of Pectoral ; (2) Pars sternalis of Pectoral ; (3) Pars
epicoracoidalis of Pectoral, (4) Pars episternalis of Deltoid ; and
(5) Pars scapularis of Deltoid. As contrasted with Rana, the
most salient difference shown in this dissection is the very large size
of the episternal head of the deltoid, which is as large as, and of
course longer than the scapular portion, and together with the
epicoracoidal head of the pectoral completely hides from view the
338 MR. F. E. BEDDARD ON THE ANATOMY [ Apr. 9,
coraco-radialis. The latter is fully exposed in Rena owing to the
slenderness of the slip, which represents in that Frog the very
large pars episternalis deltoidei of Megalophrys.
The Coraco-brachialis brevis seems te be exactly as in Rana.
The coraco-brachialis longus is a double muscle, exposed only by
cutting the pars sternalis of the pectoral.
I could not find the pectoro-cutaneous muscles which are so
conspicuous in the large Rana guppyt.
Rectus abdominis.—This muscle arises precisely as in Rana
from the pelvis, and has here the same conical form, expanding as
it does rapidly from behind forwards. Between the two Recti
lies the anterior abdominal vein, which is evident until it dips
beneath the backwardly prolonged sternum in front. The rectus
abdominis has only four Jiscriptiones tendinee. Three of these
lie behind the posterior end of the sternum. The fourth les
beneath the expanded xiphisternum, covered by it, that is to say,
when the frog is viewed from the ventral aspect. In Rana
temporaria there are five of these transverse septa, between the
several sections of the muscle. Anteriorly to the first inscriptio
tendinea the main mass of the muscle passes without a break, as
in Rana, mto the sterno-hyoid. A portion of the rectus muscle,
however, just anterior to the origin of the pectoralis abdominalis
and largely covered by the obliquus, is separable by a distinct gap
and hes at the outside of the main mass of the rectus. This
muscular slp ends in a strong tendon which is attached to the
rhomboid swelling of the sternum and seems even to reach the
coracoid beyond it.
The Sterno-hyoideus is rather more complicated than in Rana.
It is first of all found as the forward extension of the rectus
abdominis. From the expanded xiphisternum—from its anterior
edge—a flat band of fibresruns forward which dip under the tendon
of the sternal insertion of the rectus and join the main mass of
the sterno-hyoid. There is a third origin of this muscle from the
concealed (dorsal) surface of the sternum. The fibres of this
muscle arise from further back along the sternum almost from
the very extremity of the expanded xiphisternum. The more
anteriorly arising fibres form a separate muscle, distinguishable
even by a lighter colour, which runs along the inside of the main
body of the sterno-hyoid. The insertion upon the body of the
hyoid is more extensive than in either Rana or Pelodytes. The
insertions of the two muscles are in contact in the middle line of
the cartilage and they extend further anteriorly.
§ Hyoid and its Musculature.
The Hyoid and its musculature ave in some ways peculiar in
Megalophrys nasuta as compared with those of other Batrachia.
On opening the body-cavity, a strong pillar of muscle is seen
anteriorly resting on the pharynx on either side. This is the
ceratohyal with the enveloping hyvglossal muscle. In Aana the
1907. | OF A FROG OF THE GENUS MEGALOPHRYS. 339
ceratohyal is a-continuation on the same plane and in the same
straight line of the rest of the hyoid. In Megalophrys this part
of the hyoid arch is bent down at nearly a right angle with the
rest of the hyoid. The close connection of the bone and its
ensheathing muscle with the pharynx led me to assume at first
an actual anatomical connection between the two. A careful
examination, however, shows that there is no such connection,
and that the hyoid with the muscle can be raised up from its
position which is in actual contact with the pharynx. It is even.
possible that pressure may be exerted upon the pharynx by the
muscle. In the second place, this hyoglossal muscle is very much
thicker and altogether stouter than the same muscle in the very
much larger frog Rana guppyt. As in Rana, the two halves of
the hyoglossal muscle fuse and become one, and this is continued
forward in close contact and on the median line of the hyoid to
the tongue.
The Submentalis has the usual positions and relations. What.
is remarkable about it in the present species is that it is actually
(and therefore @ fortiori relatively) larger than the same muscle
in Rana guppy.
The Petrohyoidei ave, asin Rana esculenta*, four in number.
As in that frog, the petrohyoideus of Megalophrys, arising also
from the body of the byoid, is larger than the three following
divisions of the muscle which spring from the bony thyrohyals.
The last of these in J/egalophrys is a large fan-shaped muscle
which is not so very much smaller than the anterior petrohyoid.
And, moreover, the three divisions of this posterior petrohyoideus
muscle are all of fair size, and not merely in contact at their
origins (which they are not in Rana esculenta), but the middle
one actually overlaps the other two. It is important to note that
in Pelodytes punctatus, according to Ridewoody, the fourth
division of the petrohyoid is absent.
The third division of the Petrohyoideus posterior arises, it may
be remarked, from the inner edge of the thyrohyal and not as the
other two from or near the outer edge of this bone, 7.e., that
facing towards the attachment of the muscles. In Rana, Gaupp
describes the muscle in question as springing from the cartilaginous
epiphysis of the thyrohyal. This is not at all the case with
Megalophrys, where the epiphysis in question, easily detached, is
a relatively long and spur-like plate of cartilage directed outwards
and at right angles to the shaft of the thyrohyal. Tbe anterior
ot the three posterior petrohyoids does not arise from the bony
thyrohyal, but from the cartilage at its Junction with the body of
the hyoid.
The Omohyoid appears to show no peculiarities as compared with
Rana.
The Geniohyoid has the usual two origins, and it is to be noted
that these are as in Pelodytes according to Ridewood, and not as
a Gann s edition of Ecker’s ‘ Anatomie des Frosches,’ Hay, p. 139.
+ P.Z.S. 1897, p. 579.
340 MR. F. E. BEDDARD ON THE ANATOMY [ Apr. 9,
Rana according to the same observer; that is to‘say, its origin
does not extend on to the tough membrane which lies between
the posterior process of the body of the hyoid and the thyro-
hyal. Nor, in Megalophrys, as again in Pelodytes but not in
Rana, is any part of the sternohyoid inserted on to this membrane.
The reason for this is clear in the case of Megalophrys. For in
that Frog the very large posterior petrohyoid completely covers
over this space and would allow of the insertion of no muscle
‘upon it.
The Subhyoideus (the posterior portion of the submaxillaris of
some authors) is rather different in the Frog which forms the
subject of the present communication from the corresponding
muscle of Rana. In the latter it is a hyoidean muscle and arises
in the extreme lateral region of the hyoid cornua. In Megalophrys,
where these cornua are absent, I traced the muscle to the wall of
the skull just behind the tympanum.
The Hyoid cartilage is not like that of either Pelodytes or Pelo-
bates, both of them allies of the present genus. Nor does it recall
that of Rana, for I can find no trace of a cartilaginous cornu
principale or hyoid arch proper ; and as I have already stated, the
subhyoideus muscle arises from the skull-wall and did not serve to
guide me to a hyoid bar.
Xenophrys monticola obviously comes nearer to my species than
any other species of the hyoid of which I can find a description.
Of this Frog the late Prof. W. K. Parker wrote* : “ I could find no
cartilage in the hyoid arch from the Hustachian opening down-
wards until I reached the hypohyal region.” The outline is very
sunilar to that of Megalophrys nasuta. There is towards the
upper end of the diverging body of the hyoid (the hypohyal of
Parker) a nick in the cartilage on the outer side. This, as I
think, represents the nick which I figure here in Megalophrys
nasuta, and to which what appear to be vestiges of the hyoid
arches or ceratohyals are attached. Behind this in Yenophrys
monticola, as in Megalophrys nasuta, the body of the hyoid forms
a bay from which arises in my species the anterior petrohyoideus.
There is no more trace in Yenophrys than in my species of an
anterior lateral process of the body of the hyoid, unless indeed
the hypohyal really represents that, and the true anterior process
of the body of the hyoid is absent. In both these Frogs, however,
the posterior lateral process of the hyoid is present, and the thyroid
bar is well-developed and ossified, but without the spur-like carti-
Jaginous epiphysis of Megalophrys nasuta.
The accompanying figure (text-fig. 97) shows the hyoid of Mega-
lophrys nasuta. ‘The body is fairly stout and shows no traces of
an anterior lateral process. The anterior, ‘‘ hypohyal,” process on
each side is a somewhat spoon-shaped piece of cartilage in the
same straight line with the rest of the body of the hyoid. It is
not bent inwards as in Pelobates and Pelodytest. Nor could I
* “The Skull in the Batrachia,” Phil. Trans. 1881, pl. 28, fig. vil.
+ P.Z.S. 1897, pl. xxxv. figs. 10, 12.
IQOn. | OF A FROG OF THE GENUS MEGALOPHRYS. 341
find any foramen in the cartilage such as occurs in Pelodytes.
‘There was a slight thinning, however, of limited extent at a point
just opposite to the divergence of the two anterior prolongations
of the body of the hyoid, and corresponding therefore to the
marked foramina shown in the two genera Pelodytes and Pelobates
by Dr. Ridewood, whose figures are referred to below, and in
Scaphiopus*. If that be so, it would seem to follow that the tract
of cartilage lying outside of this is really the remains of the
hyoidean cornu, and the apparent solidity of the body of the hyoid
due to the complete and secondary fusion of this hyoidean cornu
with the body of the hyoid. The weak place in the body of the
hyoid is, however, so little marked that I hesitate to give to it
this morphological importance, and, moreover, it is covered by
Text-fig. 97.
Hyoid of Megalophrys nasuta.
ep. Cartilaginous epiphysis. 7. Ligament representing anterior cornua.
the attachment of the sterno-hyoideus muscle which lies to the
inside of the foramen in Pelodytes punctatus. More likely to
correspond to traces of the otherwise missing hyoidean cornu is, I
think, a stout ligament shown in my figure (text-fig. 97) which is
attached to a slight indentation near to the anterior end of the
anterior prolongation of the body of the hyoid. The position, it will
be observed, is by no means unsuitable to such an interpretation
of its nature. Nor would thisargument be necessarily at variance
with the supposition that the thinning on the body of the hyoid
in this frog is the equivalent of the foramen in Pelodytes and
* Boulenger, P. Z. 8. 1899, p. 792.
342 MR, F,. E. BEDDARD ON THE ANATOMY (Apr. 9,
Pelobates. 1 could find no splint-bone, such as Ridewood has
figured in Pelodytes as lying upon the posterior edge of the body of
the hyoid among the fibres of the sterno-hyoid muscle whose in-
sertion reaches quite to this edge. The body of the hyoid contained
no ossified tracts such as are sometimes met with in this cartilage
in other Frogs. The strongly ossified thyrohyals he, as already
mentioned, at a considerable angle with the hody of the hyoid.
The cartilaginous epiphysis of the same is not a cap at the free
end, but is attached to the outer edge and is somewhat curved.
It suggests a separate but rudimentary upper element of this
branchial arch—if it really be one, and not a mere process of the
body of the hyoid. Dr. Ridewood quotes Mr. Boulenger’s opinion
‘that the hyoidean cornua were disjointed in all those genera
which he includes in the family Pelobatide.”* My own results do
not enable me to support fully this assertion. I would rather say
that among the Pelobatidee the hyoidean cornua are clearly on the
road to disappearance. Dr. Ridewood has pointed out several
points of likeness between these structures in the Aglossa and in
the Pelobatide. I may add that the disappearance of the anterior
cornua in Megalophrys nasute and Xenophrys monticola is also
to be observed in Pipa.
§ Muscles of Shoulder and Arm.
The muscles arising from or inserted into the scapula appear
to be the same in Megalophrys as in Rana; for I identified the
following, viz.:— Dorsalis scapule, Rhomboideus anterior, Kh. pos-
terior, Levator scapule superior, L. s, inferior, Cucullaris, Serratus
superior, S. medius, S. inferior, Interscapularis, and Omohyordeus ;
besides others of which a further account seems to be desirable.
There are two of these, viz., the deltoid, and a muscle of which I
find no account in Gaupp’s edition of Hecker, and which I term
scapulo-humeralis. 1 have already referred to the pars episternalis
of the deltoid. The only further point to be remarked upon is
the apparently total absence of a clavicular head, which head I
found to be very conspicuous in the large Rana guppy.
The scapulo-humeralis is a small slender muscle running from
the scapula to the neck of the humerus, between the scapular and
adjoining humeral heads of the anconeus. I found this muscle in
Rana guppy.
§ Muscles of Thigh.
The thigh-muscles of Megalophrys are in many respects different
from those of Rana.
The most salient difference is seen on the inside of the thigh
when the skin is removed, and concerns the Semitendinosus. ‘This
muscle is then evident without further dissection, and is not at
all covered by the two Graciles. The tendon of insertion of these
two latter muscles in fact, instead of passing over the tendon of
* Journ. Linn. Soe. vol. xxvi. p. 53.
1307 OF A FROG OF THE GENUS MEGALOPHRYS. 343,
the semitendinosus (that is of course when the thigh is inspected
from the inside), passes below it. The arrangement is thus the
precise reverse of that which characterises Rana.
The Semitendinosus is then quite a superficial muscle on the
inside aspect of the thigh, visible for practically its whole extent
without dissection. In Mana it is covered not only by the graciles
but by the adductor. The semitendinosus may be termed a
two-headed muscle, but there are traces of further subdivision at.
the origin from the symphysis. ‘The lower (posterior) half of the
muscle arises by two closely contiguous heads and is to some
extent overlapped by the gracilis major. It ends at about the
end of the second third of the femu: in a stout tendon which is
continuous with the insertion on to the inside of the leg below
the knee. The anterior half of the muscle is inserted on to this
tendon soon after it is established. There is no common fleshy
origin with any part of the adductor mass such as occurs in Rana.
Unless the anterior head of the semitendinosus is to be
regarded as its equivalent, there is no Sartorius muscle. It 1s
important to notice that in the absence of a sartorius and in the
superficial semitendinosus, J/egalophrys not merely differs from
Rana but agrees with Pipa*.
§ The Lungs and the “ Diaphragm.”
The Zungs of this species are not much less in size than those
of the huge female Rana guppyi. The suspensory ligaments
appear, however, to be rather different and morecomplex. I shall
give some account of them rather asa basis for future comparison,
since but little seems to be known concerning these and other
peritoneal folds among the Batrachia. Anteriorly the right lung
is not in actual contact with the right lobe of the liver ; ‘there is
a definite and rather broad pulmo- hepatic ligament, such as is
pain in Rana guppy. This hgament runs back and at the end
of the liver-lobe becomes continuous with the postcaval vein.
It is more extensive than as well as rather different from the
corresponding ligament in Rana guppyi, and, as will be seen
presently, extends much further along the lung. This pulmo-
hepatic and afterwards pulmo-caval ligament is attached to the
edge of the lung mesiad. It does not end posteriorly in a
crescentic free edge as such ligaments often do. It runs as far
as the beginning of the kidney and then bends round and
becomes counibinomiome with a ligament attaching the dorsal surface
of the lung to the oviducal ligament and so to the dorsal body-
wall. There is thus a continuous U-shaped line of attachment,
and this traverses the lung nearly from end to end. There is
thus a similarity to the conditions observable in most Lizards
where there is both a pulmo- hepatic and a dorsal pulmonary
ligament. “It may be noted,” observed Mr. Butler, who first
contrasted f the Teiidee, which do not possess the pulmo-hepatic,
* P.Z.S. 1895, p. 838. + PB. Z.S. 1889, p. 465 footnote.
344 MR. F. E. BEDDARD ON THE ANATOMY [Apr. 9,
with other Lizards which do, “that two similar types occur
among the Amphibia. Thus in the Salamander all the men
branous attachments of the lungs and liver seem to be pr Seely
similar to those in the common Lacertilian type (e.g. Lacerta).
But in the Frog the two lungs hang freely suspended on either
side, as in the Teide.” This is clearly not true of MMegalophrys
nasuta. But it is very nearly true of Rana guppyi, but not
quite true.
In the latter Frog in fact the pulmonary ligaments, which are
illustrated in the accompanying drawing (text-fig. 99, p. 349), have
the following disposition and attachments. I have examined three
examples of this species and all of them females, and the structure
of the parts in question only differs slightly from individual to
individual. By far the greater part of the right lung hangs freely
in the body-cavity, without any attachments to neighbouring
viscera or to the parietes. The difference in this particular between
Rana and Megalophrys is quite striking. Towards the root of the
lung a ligament runs obliquely across that viseus; it is at first
atieeed to the right lobe of the liver and runs on to the post-
caval vein. The arrangement in fact is so far exactly as in
Megalophrys. But in the latter the line of attachment of this
ligament is along the longitudinal axis of the lung, whereas in
Megalophrs ys it iS more across and at right angles. ‘to this axis,
Otherwise, and save in the limited extent of the ligament, the two
agree essentially in the mode of suspension of ‘the right lung
within the body- cavity. The ligament curves down in the same
way as in Weg galophry ‘ys, and is connected with the suspensory
ligament of the oviduct. The left lung shows the same relative
differences. In Megalophrys it hasa very long line of attachment
to the stomach, while this is not the case with Rana. In view,
therefore, of these differences, it is important to compare this
genus with its undoubted ally Pelobates. In the latter Frog the
ligament attached to the lung extends for more than half-
way down that viscus in its contracted condition, and reaches as
far back as the liver in its opposite attachment. Thus there is
no doubt the Frog which [I describe here is more nearly allied in
this particular also to Pelobates thanitisto Rana. And there are
other likenesses between these genera in addition to those already
made use of by systematists.
In the case of Pipa and eaous, the anatomy of which has been
recently studied by myself *, and ten years later by Dr. Keith 7,
there are muscles which attach the lungs to the parietes. A
portion of this complex “diaphragm” is also to be found in the
Common Frog, where it consists of a sheet of muscle, a part of the
transversalis which ends upon the cesophagus and the roots of the
lungs. This has been figured in several places, and the most recent
figure known to me is in the latest edition of Ecker’s great work
* Pipa in P.Z.8. 1895, p. 827, and Xenopus, ibid. p. 841. f
+ “The Nature of the Mammahan Diaphragm &c.,” J. Anat. Phys. xxxix. 1905,
p- 253.
1907. | OF A FROG OF THE GENUS MEGALOPHRYS. 345
upon the anatomy of the Frog*. In this figure the muscle is
seen when the frog is examined as lying upon the dorsal surface
to rise gradually from the muscular parietes and to end upon the
cesophagus near to its dorsal side. It is a strap-shaped, quite flat
band of muscle.
In Rana tigrina, where I have examined the muscle carefully,
and which is a larger species and thus lends itself more success-
fully to such an examination, the muscle is very obvious and
precisely as in &, esculenta. Its fibres can easily be seen to end
upon the cesophagus. J could not, however, detect the fact—if it
be a fact—that some of these fibres cross the cesophagus to be
inserted upon the dorsal face of the lung (or rather, of course, its
peritoneal sheath). On the other hand, a portion of the muscle
in question is inserted upon the cer vical aponeurosis, which lies
in front of the lungs and heart and shuts off the neck from the
trunk. ‘These muscle-fibres pass into the aponeurosis, and may
well be attached to the lung ventrally, since they appear to reach
the pericardium beyond it.
_ The large Rana guppyt shows an interesting deviation from
these diaphragmatic structures in Rana tigrina, which I do not
think is entirely a matter of greater clearness of vi isibility owing
to its much larger size. It is, as I believe, to be put down rather
to the greater complexity i in the detailed structure of animals of a
large size as compared with their smaller allies. In this Frog the
franeversalis muscle is precisely as in Rana tigrina, save or the
fact that two considerable bands of fibres at no great distance
from each other raise themselves from the common mass of the
transversalis and pass just under the oviduct to the aponeurosis of
the lung. It seems to me to be plain that these detached slips
correspond to that portion of the corresponding muscle (with a
different origin however) which in Pipa and Aenopus are very
conspicuously attached to the lung, as is shown in the figures of
Dr. Keith + and myself %. Except im size and importance there
is thus less difference between Rana (at any rate so far as concerns
R. guppyr) and Xenopus and Pipa as regards the insertion of the
“spinal segment of the diaphragm” than might have been
inferred. As has been pointed out by Dr. Keith, “the dorsal part
of the “ diaphragm ” in Yenopus has migrated back § —so at least,
it would seem—to the ilium from hte it arises, and in Pipa
further back still, z.e. to the femur, whence the muscle in question
springs in that genus. I have, however, now to record some new
facts which tend to throw a doubt upon this interpretation of
Dr. Keith.
In Megalophrys the transversalis sheet to the esophagus is a
muscle of very great extent ; it 1s, for instance, fully twice the
length (not relatively, but actually) of the same muscle in Rana
* © Anatomie des Frosches,’ by Gaupp.
ii Loe. cit. fig. 10, p. 254, & fig. 11, p. 255
di Hoc: cit. fig. 1, p. 842.
§ These facts were also correctly noted by mysclf.
346 MR. F, E. BEDDARD ON THE ANATOMY [ Apr. 9,
guppyi. It follows, therefore, that as the general build of these
two Frogs, i. e. the proportions of the skeleton, is much the
same, the origin of the muscle in Megalophrys must differ from
that of Rana. And, as a matter of fact, it does. In Megalophrys
the muscle runs back into quite the pelvic region, and its posterior
part arises actually from the ilium. In respect of this muscle,
therefore, Wegalophrys offers us actually what is a theoretically
possible condition in the ancestor of both Rana and Xenopus. Of
a continuous sheet of muscle such as occurs in Megalophrys, the
anterior part only has been retained in Rana and the posterior
Text-fig. 98.
The cesophageal sheet of transversalis muscle in Megalophrys nasuta.
al. Hsophagus. Cocc.I1. Coceygeo-iliacus muscle. h.g. Hyoglossus. 0.d. Oviduct.
cs. (Ssophageal muscle.
part only in Xenopus. A careful examination of this muscle in
Megalophrys has convinced me that no fibres escape from it on to
the lung or any other viscera. Its attachment to the cesophagus,
and to the cesophagus only, can be easily followed. The oviducal
membrane which it underlies is clear and transparent throughout,
with no muscular fibres traversing it.
Tn view of the interesting differences which the transversalis
muscle of Megalophrys shows to the corresponding muscle of the
1907.] OF A FROG OF THE GENUS MEGALOPHRYS. 347
other genera of Batrachia described above, I am particularly glad
to have had the opportunity of dissecting, and to be able to record
the appearance of, the muscles of this region of the “ thorax ” in
Pelobates fuscus; for this genus is referred by Boulenger to
the same family, Pelobatidee, as that which contains Meg galophry Ys.
That Boulenger’s view is more correct than that of Mivart * and
some others, 1s testified to by the structures of these various
muscular arrangements which have been considered in the
foregoing pages. The figure (text-fig. 95, p. 334) is a drawing
of the interior of the body of a male WPalhaies Juscus, with such
viscera removed as interfere with the proper view of the trans-
versalis muscle. It will be noted that that muscle is very
extensively developed and that its origin reaches back not only
to the ilium, but for some way along that bone, where it overlaps,
but hardly conceals, the coccygeo-iliacus.
The whole of the transversalis muscle in Pelobates is obviously
fixed to the cesophagus, in which there is a slight difference from
Megalophrys, where the hinder end of the muscle does not appear
to reach the cesophagus but to terminate upon the oviducal
membrane. The Pelobates however, as | have mentioned, was a
male; but the corresponding part of the muscle were it present
would naturally lie upon the median mesentery. The muscle is,
moreover, at first sight rather reduced in size as compared with
that of Megalophrys. But a close study shows that this is due to
the semilunar excavation of the posterior margin of the same,—
that really its origin extends quite as far back as in Jegalophrys.
Moreover, in both of these Frogs the muscle completely covers
and conceals the ilio-costal muscles; anteriorly a state of affairs
which is not found in Rana &c., as is depicted in the accompanying
figure (text-fig. 98). In Pelobates the distinction between this
muscle and that which is applied to the terminal cervical aponeu-
rosis is very marked; their fibres do not show any continuity
anywhere. This muscle—the ventral part of the diaphragm, as
Dr. Keith terms it—1is better developed in Pelobates than in Rana.
There is hardly any membranous portion left. This is plainly
shown in the figure.
I have also examined into the condition of this cesophageal
sheet of the transversalis in other genera. In the South American
Ceratophrys ornata the muscle is easy to see and is disposed rather
differently from the muscle in the genera that have been already
referred to. It is not extensive as in Weqalophrys ; but is, as in
Rana, limited to the fore part of the visceral cavity. Instead of
forming at its origin at any rate a single continuous sheet of
muscle, it is in Ceratophrys distinctly divided into three portions.
The present 1s one of the species of the genus which possesses a
large dorsal ossification in the cutis, which is firmly articulated
to the vertebral transverse process, from the surface of which the
muscles in question arise. ‘The innermost or posterior of the three
* “The Classification of the Anurous Batrachians,” P. Z.S. 1869, p. 290 &e.
348 MR. F. E, BEDDARD ON THE ANATOMY [ Apr. 9,
slips (on the right side of the body) is the one which is attached
to the cesophagus. It is, as already said, short as in Rana, and
does not extend far back in the body. The middle slip partly
covers this and is of considerably less size, that is to say, breadth,
for the length hardly differs. It lies at a different plane from the
first slip of the entire muscle, being more ventrally directed in its
course. It is attached to the membrane bearing the oviduct,
which is in its turn connected with the membrane at the root of
the lung. The outermost portion of the muscle is the largest and
is chiefly spread over the cervical aponeurosis, a good many of its
fibres being directed towards the lung. On the left side of the
body of the example of this species which I dissected, there was
not quite so marked a distinction between the first two parts of
the muscle. There is thus in this species of Frog a very much
more marked subdivision of the muscle than exists in Rana guppy,
which is perhaps due to, and is at least connected with, the fact
that the origin of the muscle in Ceratophrys is more spread over
the surface of the transverse process, which is moreover enlarged
and rather firmly connected with the dorsal scute. This transverse
process extends outward far beyond the transverse processes of the
vertebrae which follow it. I am at present unable to go into
detail; but a dissection of Bufo agua and of Rhacophorus* sp.,
has shown that in these species also there is some specialisation of
the slips of muscle passing to the cesophagus and lung.
§ Alimentary Canal.
The alimentary canal and the appended glands of Megalophrys
nasuta show certain small variations from the structure of those
organs in Rana esculenta and RF. tigrina. But the conditions
obtaining in Rana guppyt (text-fig. 99) prove that the form of the
liver, for example, is to be used with caution as a generic character,
as do also the variations in the form of this organ in the Common
Frog. In Rana guppy, for instance, the two lobes of the liver
represent the entire’ subdivision of that organ. The right and
left lobes were not further subdivided in any ‘of the three examples
at my disposal. And it may be remarked that three examples
selected at random are a very fair test of a characteristic. More-
over, in all of these three specimens the right and left lobes were
of approximately the same size. They were connected below the
cesophagus by a narrow isthmus of hepatic tissue. Now in the
Common Frogs of this country the left lobe is decidedly the larger
of the two. In the example of Rana tigrina which I dissected
the outer of the two lobes of the left division of the liver was,
though only slightly, the larger; it is the same lobe which is figured
as the larger in Rana esculenta. Megalophrys agrees with Rana
esculenta and 2. tigrina as to the general disposition of the liver-
* From Borneo. I owe this specimen also to the kindness of Dr. Charles Hose.
+ Cf. e.g.,‘The Anatomy of the Frog’ by Ecker, Engl. trans. by Haslam: Oxford,
1889, p. 295, fig. 194.
1907.] OF A FROG OF THE GENUS MEGALOPHRYS. 349
lobes. The left is very much the larger of the two, and it is sub-
divided into two lobes of unequal size of which the outer, situated
more ventrally, is the larger *. In Pelobates fuscus the right half
of the liver is the smaller; the left lobe is also (as in Megalophrys)
composed of two very distinct lobes of approximately equal size, the
inner however being in this case rather the larger. The pancreas
of both Megalophrys and Pelobates differs from that of Kana as
Text-fig. 99.
Liver and adjacent viscera of Rana guppy?.
g.b. Gall-bladder. 2. Liver. Zw. Lung. m. Mesentery attaching liver to lung &e.
figured by many authors. Its lowest portion is massive in these
two genera; but, whereas in Pelobates this region of the pancreas
is firmly attached to the stomach, it is attached to the duodenum
in Megalophrys. The separation of the stomach from the duodenum
* Duméril & Bibron, ‘ Erpétologie Générale, t. vii. p. 456, remark also (of
M. montana) that the right lobe is but slightly developed and that the left is divided
into two.
Proc. Zoou. Soc.—1907, No. XXIV. 24
350 MR. F. E. BEDDARD ON THE ANATOMY [| Apr. 9,
is more marked in Megalophrys than in either Pelobates or Rana.
The pylorus is situated a little way beyond the lowest point
of the U formed by the stomach and the duodenum and is
marked externally by a raised annular fold.
$ Ovaries and Oviducts.
The whole abdominal cavity of this Frog was filled with eggs
which were not contained within the oviducts, but in the ovaries
themselves. The lobes of the ovary packed tightly with ripe ova,
among which were immature ova, were polygonal through mutual
Text-fig. 100.
Oviduct of Megalophrys nasuta.
o.d. Dilated lower part of oviduct. ov. Ovary. m. Muscular band attaching
ovary to cecum of oviduct.
pressure. The mesovarinm which joins the mesocolon is remark-
able for the fact that posteriorly its margin is strengthened by a
thick and strong band of muscular fibres. I could not observe
any structure of this kind in Rana guppyi in the same region.
This band of muscles in Wegalophrys is shown in the accompanying
figure (text-fig. 100). It will be there seen that it is broken up
into wider and narrower strands of muscle, which are partly con-
1907.] OF A FROG OF THE GENUS MEGALOPHRYS. 301
tained within or lie upon the mesovarium and thus run towards
the median dorsal line of the body, and are partly imserted upon
the distal dilated sac in which the oviduct ends (as in Ranw &c.),
and by which it opens into the cloaca.
Traced in the opposite direction, the fibres soon lose themselves
upon the ovary. The strand of muscles which is inserted upon
the terminal chamber of the oviduct is much the thickest. It
was, however, not absolutely accurate to speak of this band of
muscular fibres as being inserted upon the dilated posterior
chamber of the oviduct. They are really inserted upon a distinct
and rather tubular diverticulum of the same, which lies to the
inside of the oviduct and naturally points towards the ovary.
This diverticulum is certainly totally absent in Rana guppy?,
where—considering the size of the Frog—it would be easy to
detect were it present. The band of muscles continuing on this
diverticulum to the ovary associates it particularly with the gonad.
The association might be physiological as well as anatomical. For
a swelling of the ovaries would tend to fall upon the diverticulum
of the uterus, possibly thus stimulating its growth and that of the
uterus at the time when it is required to be large for the reception
of the ova.
On the left side of the body the structure of the various bands
of muscle and the diverticulum of the uterus was not absolutely
identical. In fact, there was no appearance of a stout band of
muscle applied to the diverticulum of the uterus, only fine fibres
in the mesovarium. Nor was the diverticulum so conspicuously
marked as on the right side. The two uteri join just before they
open into the cloaca on a marked papilla by a single orifice, as in
some genera.
The internal orifice of the oviduct in this Frog has a position
which differs rather from that of the oviducal aperture of Rana
guppyi. n the latter it is distinctly to the outside of the lung;
in my species it is situated as distinctly to the inside of the lung
base. In both cases this funnel-shaped expansion lies upon the
ligament attaching the liver to the anterior wall of the abdominal
cavity.
§ Summary of Principal New Facts.
Tt will be convenient to briefly summarise the chief new facts
contained in the preceding pages. They are as follows :—
(1) The genera of the family Pelobatidee should perhaps be
increased by the addition of one for a Bornean species, possibly
identical with Megalophrys nasuta (auct.), which differs from all
other genera of that family in certain features.
(2) This species agrees with Pelobates (¢as to other Pelobatidw)
and differs from Rana, Ceratophrys, and some other genera, in the
great extent of the visceral layer of the transversalis muscle
(‘diaphragm ”), which is attached along a greater length of vhe
pharynx and arises from the transverse processes of several ver-
tebre and from the ilium for a long way down, covering over ana
24*
352 MR. W. P. PYCRAFT—CONTRIBUTIONS | Apr. 9,
completely concealing the ilio-lumbaris muscle, which is entirely
exposed in the other genera mentioned.
(3) The equivalent muscle of Ceratophrys, though arising as In
Rana and other genera from the transverse process of the fourth
vertebra only, is specialised into three separate slips, supplying
respectively the pharynx, lung, and cervical aponeurosis. This
specialisation of the muscle is also be seen in Rhachophorus and
Rana guppy.
(4) In Megalophrys nasuta the sacral vertebra is single and
fused with the coccyx and the vertebre are proccelous.
(5) The hyoid in this Frog agrees with that of the Pelobatidze
generally in the deficient development of the anterior cornua.
These are only represented by a ligament, and the hyoid as a whole
most closely resembles that of Xenophrys monticola as figured by
W.K. Parker. The musculature of the hyoid does not materially |
differ from that of Rana.
(6) The musculature of the back shows characteristic differences
among different genera of Batrachians. The genera of Ranide
and Cystignathide and Bufonide examined show fewer differences
among themselves than any of them do from the Pelobatide,
Pelobates and Megalophrys nasuta.
(7) In the last two is a peculiar muscle apparently belonging
to the ilio-lumbar complex, running from the middle of the ium
directly to the transverse process of the second vertebra, which
may perhaps be the equivalent of the “ Musculus pulmonum pro-
prius” of Pipa. Other comparisons between the muscular system
of Megalophrys nasuta ave remarked upon in the present paper ;
and other authors have commented upon likenesses between the
Aglossa and the Pelobatide.
4. Contributions to the Osteology of Birds.—Part IX.
Tyrannt ; Hirundines ; Muscicape, Lani, and Gymno-
rhines. By W. P. Pycrart, F.Z.8., A.L.S., &e.
[Received March 18, 1907. |
(Text-figures 101-1 04.)
ConTENTS.
i. Introductory Remarks, p. 352. vi. The Pelvic Girdle, p. 369.
ii. The Skull of the Adult, p. 353. vil. The Pectoral Limb, p. 370.
ii. The Vertebral Column, p. 366. vill. The Pelvic Limb, p. 371.
iv. The Ribs, p. 367. ix. Summary, p. 372.
y. The Sternum and Shoulder-Girdle, x. Key to the Osteological Characters
p. 367. of the Skull, p. 378.
i. IyrropuctoryY REMARKS.
Although it cannot be claimed for the following pages that
they contain a complete account of the osteology of the several
groups enumerated above, they may at least be said to contain
1907. | TO THE OSTEOLOGY OF BIRDS. 353
the most exhaustive account that has yet been written. Anything
like completion will be impossible for very many years to come,
owing, largely, to the difficulty in obtaining material. Large as
is the British Museum collection of skeletons of these birds,
there are considerable gaps to fill.
As a contribution towards our knowledge of the Passeres this
paper will, I trust, prove useful. I have endeavoured especially
to throw light on the perplexing questions of classification, but
with what measure of success time must prove.
lil. THE SKULL OF THE ADULT,
The Occipital Region.
The plane of the occipital foramen is tilted backwards, owing
to the relatively large size of the brain-case, so as to lie almost
horizontally : that is to say, almost in a line with the long axis of
the skull.
The supraforaminal ridge is not very well marked, in so far as
its extension laterad of the foramen is concerned. It generally
terminates on the bulla which forms the outer wall of the pos-
terior tympanic recess.
A pair of small foramina may be mentioned here which admit
branches of the vena occipitalis. In Pitta, Phytotoma, Pitangus,
Tyrannus, Prionops, Huryceros, and Lanius, for example, these
foramina will be found forming the termination of a pair of
well-marked grooves traversing the supraoccipital immediately
above the occipital foramen ; and extending upwards towards the
middle line to pass into the foramina in question on either side
of the occipital crest. But in Artamia, Artamus, Corvinella, and
Tylas, for instance, these foramina seem to have shifted, inasmuch
as they appear much lower down—immediately above the rim of
the occipital foramen ; while in Gymnorhina, Pityriasis, Dicrurus,
and Vireolanius they are found on the free edge of the rim itself.
A cerebellar dome is nowhere very well developed: it is
practically absent except among the Tyrannide and Artamide.
The lambdoidal ridge is generally moderately well developed ;
though in no case is it so strongly marked as in Pitta among
the Tyranni, and in Lalage and Graucalus representing the
Campophagide ; it forms, however, only a very inconspicuous
feature of this region of the skull.
The Cranial Roof (text-fig. 101)—Save only in the Tyrannide,
the cerebral dome is broad and well-rounded, while the contour
of the cerebrum is plainly indicated, and the interorbital region
of the frontal is wide. In this last character it would seem that
the Campophagide, Prionopide, and Laniinz differ, in that this
region is narrow. Puaradisea is peculiar in having a relatively
long and narrow cranium, but with the cerebral dome well
marked. In the Tyrannide the cerebral dome is conspicuously
depressed, but wide; the interorbital region is generally wide in
354 MR. W. P. PYCRAFT—CONTRIBUTIONS [Apr. 9,
this family, but in a few genera the reverse is the case, e. g.
Myiarchus arundinicola.
No members of the groups here dealt with have developed a
true nasal-hinge, but this is feebly represented among certain
Tyrannide, e. g. Megarhynchus, Rhynchocyclus, and Contopus.
The Base of the Skult.
Basipterygoid processes are not even represented by vestiges.
The basitemporal plate exhibits the usual triangular form, the
base of which is bounded by a more or less well-marked pre-
condylar fossa ; while its two sides have fused, for the greater part
of their length from the base forwards, with the ossified con-
nective tissue forming the anterior wall of the recessus tym-
panicus anterior—converting the HKustachian grooves into tubes
opening at the apex of the triangle. The posterior angles of this
plate contribute to form the inferior margin of the tympanic
recess. Although this plate presents certain differences, when a
large number of genera belonging to different families are
examined, these differences are of too subtle a character to be of
any use for systematic purposes.
The Lateral Aspect of the Cranium.
The tympanic cavity, as seen in the dried skull, may be described
as moderately well developed. In some cases, however, a much
more spacious cavity is formed by the extension of the lateral
wing of the exoccipital. As a rule, this aperture is oval in shape,
but in some cases, e. g. Phytotoma, Vireolanius, Lalage, and
Prionops, it forms a vertical slit, the distance from the shaft of
the quadrate in front to the lateral exoccipital wing behind
being much less than the height of the aperture, whereas,
generally, the vertical axis is but little longer than the horizontal.
More rarely the free edge of the lateral exoccipital wing is
turned sharply forwards and somewhat inwards, so that a large
and nearly closed resonant chamber is formed having a sub-
crescentic external aperture, as in Paradisea for example. In
the Gymnorhine and in Artamia, one of the “ Vangide,” the
free edge of the lateral occipital wing, instead of sweeping
upwards and forwards so as to pass imperceptibly into the
processus articularis squamosi, is continued upwards till it bisects
a ridge running from the squamosal head of the quadrate across
the shallow temporal fossa or along its inferior border. This
peculiarity will probably be found to obtain in other allied
genera, while it occurs also in some Synallaxine.
Within this cavity there will be found a strong buttress of
bony trabecule for the inner head of the quadrate, in addition to
the usual orifices marking the apertures of the anterior, superior,
and posterior tympanic recesses, and the fenestre rotunda and
ovalis. The superior and posterior tympanic recesses, it may be
remarked, are relatively small: in Artamia they can hardly be
traced while in Laniarius, for example, they attain their maximum
1907.4 TO THE OSTEOLOGY OF BIRDS. 355
development. Opening immediately above the squamosal and otic
articular surfaces for the quadrate, the latter leads upwards intoa
chamber filled with cancellated tissue.
Text-fig. 101.
Lateral aspects of the skulls of :—
a. Gymnorhina organicum. b. Laniarius poliocephalus. @. Sayornis cineracea.
am. = anterior nares; a.0.p. = antorbital process; 7. = lachrymal; pa. = palatine;
p.o.p. = postorbital process; pt. = pterygoid; p.z.s. = processus zygomaticus
squamosi.
The Postorbital Region presents one or two noteworthy |
characters. The Pittide alone develop extensive, but shallow
temporal fossee, which are separated one from the other, in the
middle line, only by a median sagittal ridge, In the Tyrannide
and in the Phytotomide this fossa is, though narrower, relatively
deeper than in the Pittide; but it is confined entirely to the
lateral aspect of the cranium, extending backwards, roughly, as
far as a point midway between the free edge of the tympanic
cavity and the centre of the lambdoidal ridge: the distance
varying in different genera. In the remaining Families dealt.
356 MR. W. P. PYCRAFT—CONTRIBUTIONS [ Apr. 9,
with in this paper, the temporal fossa terminates immediately
above the tympanic cavity: excepting the Muscicapide, where it
is either extremely reduced or absent.
The postorbital process is obsolete in Phytotoma—another of
several features which this Family shares with the Pittide. In
the Tyrannide it is only very feebly developed; and the same is
true of the Hirundinide, Muscicapidee, Campophagide, and the
Lanune at least among the Laniide.
The reduction of this process is due to specialisation: and in
the forms here dealt with, it has been accompanied by an equally
gradual reduction of the “temporal fossa.” In all these cases,
the process has travelled downwards, nearer and nearer to the
processus zygomaticus squamosi, and this is nowhere better shown
than in a comparison of the skulls of MWuscicapa and Terpsiphone
for example; since in the latter the two processes are almost
confluent, and further, by the obliteration of the ‘‘ temporal
fossa” the superior margin of the tympanic cavity has extended
forwards to join the postorbital process, In the Muscicapidee this
margin has crept forwards no farther than the hinder border of
the remnant of the fossa. In the Laniine, the reduction of the
postorbital process has been accompanied by a corresponding
increase in the size of the processus zygomaticus squamosi, which
has, in consonance with the use of the jaws for tearing prey,
shifted forwards so as to lie midway between the squamosal
articular surface of the quadrate and the postorbital process.
Among the Laniiform-types, the Gymnorhide and Malaconotide
have the postorbital processes of large size, and so also have the
Dicruride, Hurycerotide, Artamide, Vangidee, and Prionopide.
The Gymnorhide, Artamide, and Vangide, by the way, are
peculiar in that the zygomatic process of the squamosal is
bifurcate : or, in other words, mesiad, and a little in advance, of
the normal process, there is found a second though slighter peg.
So far I have not met with this in any other families.
The orbits present no very important characters in this
connection. Only in the Gymnorhide and Vangide, it would
seem, is the orbito-sphenoid ossified, or the interorbital septum
perforated by but a single small fenestra. All the remaining
genera herein dealt with have the orbito-sphenoid but slightly, if
at all ossified, and the interorbital septum pierced, either by a
long narrow upper slit and a wider, inferior fenestra, or by one
large fenestra absorbing almost the whole septum as in Paradisea.
The Ethmoidal Region.— With that portion of the mesethmoid
which takes part in the formation of the interorbital septum I
have just dealt. The lateral mesethmoidal wings of the antorbital
plates—called by some pre-frontals-—-which bound the orbit
anteriorly, present but few features of special note. Gymnorhina
is somewhat remarkable in this respect Inasmuch as these plates
extend outwards and upwards, one on either side, to form a pair of
wings of considerable size, and projecting laterally on either side
of the nasals, thus greatly increasing the depth of the orbital
chamber.
19072] TO THE OSTEOLOGY OF BIRDS. 307
The lachrymal, which is attached, as a rule, to the outer margin
of the antorbital plate, demands some notice. Unfortunately, in
a very large proportion of prepared skeletons of birds this is
absent, often because of its reduced size which causes it to be
overlooked by the macerator. And thus it becomes impossible to
say, when this ossicle is missing, whether it has been lost during
maceration or suppressed. That suppression has taken place in
not a few instances is almost certain, inasmuch as in skeletons in
this collection it may be found reduced to the vanishing point.
In the Families which form the subject of this paper, the
lachrymal is least reduced in Graucalws among the Campo-
phagidee, and in Laniarius among the Laniide. In Graucalus it
is roughly D-shaped. and attached to the anterior face of the
antorbital plate; the free ends of this horseshoe curve are
directed outwards, the lower limb projecting beyond the outer
border of the plate, and resting on the quadrato-jugal bar. It is
formed of highly pneumatic bony tissue and has a conspicuously
inflated appearance. In Laniarius this ossicle is almost rod-
shaped, and has its inferior, free end twisted outwards, con-
siderably swollen, and resting on the quadrato-jugal bar. The
main shaft is attached to the anterior face of the antorbital plate,
while its upper end is wedged in between the nasal bone and the
antorbital plate. Gymnorhina agrees very nearly with Laniarius
in this respect, differing chiefly in that the inferior end of the
ossicle rests In a facet on the quadrato-jugal bar. In Artamus
and Artamia it is greatly reduced—ain the former, for example,
being represented only by a very delicate rod, with a cordiform
inferior extremity which does not reach the quadrato-jugal bar, but
which projects slightly beyond the level of the antorbital plate.
Prionops in this matter resembles Artwmus, except that the rod is
somewhat thicker and not markedly inflated at its inferior end.
In Huryceros it has the form of a sigmoidally curved rod attached
to the face of the antorbital plate, as in the preceding cases, and
extending downwards to articulate with a tumid prominence on
the quadrato-jugal bar. In the Tyrannide it is attached, not to
the anterior face, but to the outer, free edge of the antorbital
plate, and extends upwards to the level of the nasal, and down-
wards to the quadrato-jugal bar. In shape it may be described
as spatulate, the blade being flattened along its mesial border
where it comes into contact with the antorbital plate; while the
handle has a strong outward curvature. Finally, in Phytotoma
all that is left of this ossicle is a minute spicule attached to the
outer edge of the antorbital plate. In no ease is an orbital
process developed ; and only in Laniarius is the shaft perforated
by a conspicuous pneumatic foramen.
The ossification of the nasal septum and of the walls of the
olfactory chamber gives this region of the skull, in the groups now
under consideration, an unusual importance.
In the Tyrannide, this septum, in its simplest form, as in
Pitangus, extends from the cranio-facial fissure to the anterior
end of the narial orifice, and has a knife-like free edge dividing
398 MR. W. P. PYCRAFT—CONTRIBUTIONS [Apr. 9,
the anterior palatal vacuity into right and left moieties. The
dorsal expanded plate, which gives the usual T-shaped form to the
mesethmoid, is here continued outwards and downwards in the
form of a small quadrate plate of bone depending from the mesial
border of the posterior region of the narial orifice. In Sayornis,
this ossification of the dorsal mesethmoidal plate extends the whole
way along the nasal process of the premaxilla instead of being con-
fined to a small area of the nasal orifice, but the lateral extension
of the plate so formed is not great, and is most noticeable near the
middle of the orifice. The median, vertical region of the septum
has its free edge expanded to form a narrow plate, grooved along
its inferior surface down the middle line, and extending nearly
the whole length of the anterior palatal vacuity, the hinder end
of the plate being forked. And this obtains also in Contopus,
Teniopterus, and Rhyncocyhclus—and doubtless in many other
genera.
The Muscicapide have a nasal septum agreeing in all respects
with that described as obtaining in Contopus and Sayornis for
example. In the Campophagide, however, this septum is either
altogether absent, in the dried skin, or is represented only by a
feebly ossified ridge running down the inferior aspect of the
nasal process of the premaxilla.
Among the Laniide, as the group stands in Dr. Sharpe's
‘Hand-list’ (8), the ZLaniine agree with the Tyrannide and
Muscicapide in having the inferior border of the nasal septum
expanded by lateral flanges, but these are, in the group now
under discussion, very narrow.
In Sphecotheres—which genus will be discussed in detail later
(p. 376)—the nasal septum is of this type. And the same is true
of the Prionopide. In this family, however, secondary ossifi-
cation of the cartilaginous outer wall of the nasal capsule conceals
the septum from the inferior aspect of the skull. At first sight,
however, the existence of this floor beneath the septum would not
be suspected; but rather it would be regarded as the septum
itself, which had developed unusually wide flanges, since it has all
the appearances of a septum such as occurs in the Tyrannide,
differmg therefrom, mainly, in being fused anteriorly with the
premaxillary palatal plate.
In the Phytotomide, and in Corvinella*, Artamide, and the
Gymnorhine among the Laniiform types, the wall of the vesti-
bulum is ossified ; and this in sucha way that the postero-superior
angle of the normal narial aperture is cut off to form a slit-like
aperture, bounded mesially by the vestibular wall, and externally
by the descending process of the nasal. Traced backwards this slit
leads into the lachrymo-nasal fossa, and thence into the choane.
The restricted narial aperture is small, and circular, owing to the
ossification of the cartilaginous nasal capsule—the vestibulum. .
It leads into a chamber, small in Artamia, large in Artamus
containing a well-developed concha media. This turbinal in
* Lanius excubitor agrees with Corvinella in this respect and differs from
the remaining members of the genus.
1907.] TO THE OSTEOLOGY OF BIRDS. 309
Artamia is scroll-shaped, and attached to the floor of the chamber
and to the horn of the bifurcate vomer.
In Artamus the chamber is much larger, and, as in Artamia, 1s
open behind. The septum nasi is perforate, placing the right and
left chambers in communication, in the dried skull. The concha
media is large, conch-shaped, and attached by its posterior end,
caudad of the tip of the vomer, to the extreme hinder end of the
vertical outer wall of the vestibule; so that this turbinal lies above
at its posterior end—by an
upward, vertical, flange developed by the palatine. Corvinella
resembles Artamia in this matter; and the same is true of
Gymnorhina, but the concha media of each side is here attached
to the tips of the bifurcate vomer, while its wall is cribriform.
The floor of the olfactory chamber, in the forms now under
discussion, demands a more detailed notice than is usually the
case with this region of the skull.
In all save the Artamide, Prionopidee, and Gymnorhide, this
floor is cartilaginous, so that, in the dried skull, there is a large
palatal foramen which is divided by a median partition—the
septum nasi. This, generally, has its free edge broadened by a
flange along each side, thereby reducing the size of the foramen.
In the exceptional cases this floor becomes completed by the
ossification of the nasal capsule in connection with the septum,
converting the palate into one of the desmognathous type. That
this desmognathism is caused by the ossification and fusion of the
inturned capsular wall of the vestibule with the inferior border of
the nasal septum, and not by the fusion of the maxillo-palatines
with the septum, is clear from a study of the palate of say Vureo-
lanius (text-fig. 102 e, p. 362), where this ossified capsule extends
backwards above the maxillo-palatines, or of the Prionopide (e. g.
Prionops), where a narrow chink is left on each side of the septum
immediately in front of the vomer. In Paradisea the septum
descends as it were, so as to close up a median chink, left between
the right and left portions of the vestibular floor, while both floor
and septum are sharply truncated posteriorly, so as to impinge on
the maxillo-palatine process and the vomer (text-fig. 103 b). Inthe
Gymnorhide the vestibular floor meets beneath the nasal septum,
but it has not blended with the palatal surface of the premaxilla,
which presents a sharply defined posterior border into which the
ends of the palatines are thrust.
The Premaxilla and Nasal.
These bones present no characteristics sufficiently remarkable
to dwell upon here. Suffice it to say, that the palatal surface of
the premaxilla is often apparently increased by fusion with the
ossified nasal capsule, while the nasals are always holorhinal; and
in some cases, as has already been pointed out, the form of the
nasal aperture has been altered by a similar ossification of this
capsule.
In the Pittidee and in Terpsiphone among the Muscicapee there
is a fairly well-developed nasal hinge.
360 MR. W. P. PYCRAFI— CONTRIBUTIONS [ Apr. 9,
The Maxillo-jugal Arch.
The maxilla, in the skulls now under analysis, presents features
of unusual interest, inasmuch as this element of the jaw retains
more primitive characters than are to be met with among the
remaining Oscinine Passeres. My examination of this bone must
begin with the Tyrannide. Though in the adult the extent of the
maxilla cannot be determined, the palatine process therof forms a
large triangular plate with its apex curved backwards in a semi-
circle, almost meeting the free, anterior, end of the vertical
descending, seroll-shaped plate of the palatine (text-fig. 102,
me.p.) and underlying the free end of the vomer.
In the Pittide this maxillo-palatine plate has perhaps preserved
even more of its original shape, since it is larger and less pointed
at its apex (fig. 102 d, ma.p.).
The Muscicapide present the early stages of transition which
eventually produced the type of maxillo-palatines met with in the
higher Passeres. In Mewtonia these processes are broad, strap-
shaped, and deeply pitted below by a pneumatic foramen, while
their direction is backwards. In Zerpsiphone the maxillo-palatine
processes are long and narrow, and spring from a broad shelf of
bone which passes forwards into the palatine process of the pre-
maxilla ; and the same is true of Piezorhynchus. In Muscieapa the
final stage is reached : the maxillo-palatine being reduced toa long,
slender, curved bar springing from a rod-like base. The free end
of the process almost touches the free end of the descending
vertical scroll of the ventral aspect of the palatine.
The Campophagide closely resemble the Muscicapide in the
form of the maxillo-palatine process ; it appears to differ chiefly in
that it is relatively shorter than in the Muscicapide.
Among the Laniide, as at present constituted, this region of
the palate presents two strongly contrasted phases of development.
In the less specialised Laniine, e. g. Lanius, Laniarius, and
Dryoscopus, the maxillo-palatines do not differ essentially from
these elements in the Campophagide, a large palatal vacuity
bounding these processes in front. But in Corvinella the an-
terior palatal vacuity is partly filled up by the ossification of
the vestibular floor. The Prionopide and the Dicruride agree
with Corvinella in this respect. The Gymnorhine, included by
Dr. Sharpe with the Laniide, differ markedly from the Laniine
‘in this matter; and this on account of the ossification of the
vestibular wall and nasal septum already referred to (p. 358).
In Pityriasis and Gymnorhina the anterior palatal vacuity is
closed, in part by the ossified floor of the vestibulum, and in part
by the nasal septum, so that only a small pit is left infront of the
vomer. The Artamide and Vangide in this matter agree with
the Gymnorhine. The palate of the aberrant rocharis
(Huryceros) in many respects recalls that: of the Gymnorhine
(p. 359) and Artamide, but it differs in that the maxillo-palatines
in front of the maxillo-palatine processes are inflated, forming a
1907. | TO THE OSTEOLOGY OF BIRDS. 361
mass of cancellated tissue, surrounded, above and below, by a large
air-sinus, so that a considerable space is left, in the dried skull,
between the ossified floor of the vestibulum to which the palatines
are attached and this cancellated mass.
The quadrato-jugal bar affords no characters requiring comment.
The Vomer, Pterygoid, and Palatine.
The vomer is of the typically Passerine type, but presents certain
features worthy of note. Broad, and of moderate length, its
ventral aspect, in the simpler forms, e. g. Tyrannus, is that of an
oblong plate of bone cleft in the middle line from behind forwards
for rather more than half its length, and sharply truncated in
front. The right and left posterior limbs pass insensibly into
the palatines. Anteriorly, the vomer expands to form a pair of
thickened quadrangular horns the external, lateral faces of which
lodge a more or less cup-shaped depression.
The Tyrannide and Muscicapide agree very closely in the form
of the vomer. ‘The Campophagide, however, would appear to be
even simpler in this respect than either of these two Families,
inasmuch as in Campophaga the vomer is anteriorly sharply
truncated and lacks the quadrangular anterior horns, though an
approximation to these is to be seen in the vomer of Jylas.
In the Laniine the vomer, as in the Tyrannide, expands,
anteriorly, into a pair of quadrangular horns, but each sends back-
wards from its antero-dorsal angle a thin plate of bone which,
running backwards along the outer margin of each side of the
dorsal aspect, gives the vomer, when seen from above, a trough-
shape. Further, in Laniarius, the quadrangular horns referred
to afford, on either side, an articular surface for the mavxillo-
palatine process.
The Dicruride differ from the typical Laniide in the form of
the vomer chiefly in that in the former it is relatively broader,
especially at its anterior end which is sharply truncated. The
increased width, anteriorly, is due to the fact that the “‘ horns”
project outwards and forwards instead of directly forwards. These
horns, furthermore, are peculiar in that they send downwards a
broad facet to articulate with the spatulate maxillo-palatine
process. The terminal “horns” of the vomer rise upwards to
embrace the free edge of the nasal septum, and are continued
backwards along each side of the dorsal aspect of the vomer,
thereby converting this into a shallow trough.
In the Gymnorhinz the vomer is more conspicuously modified
than in any other of the types herein discussed; being short,
antero-posteriorly, and having a well-defined median ventral keel
and deeply trough-shaped dorsal aspect. These characters are
perhaps most marked in Gymnorhina, though in Pityriasis they
are scarcely less so. in the former genus the free end (anterior)
of the vomer is deeply cleft, while the under surfaces of the
two resultant horns are cut away, or hollowed out, so as to
362 MR. W. P. PYCRAFIT—CONTRIBUTIONS evar. 2)
Text-fig. 102.
Ventral aspects of the skulls of :—
a. Artamus leucogaster. b. Sayornis cineracea.
e. Terpsiphone. d. Pitta baudi.
e. Vireolanius leucotis. f. Vireo olivacea.
@.0.p. = antorbital process; mx.p. = maxillo-palatine process ; 7.s. = nasal septum ;
pa. = palatine; pt. = pterygoid; ¢.=quadrate; ve. = vomer.
1907. } TO THE OSTEOLOGY OF BIRDS. 363
Text-fig. 103.
Ventral aspects of the skulls of :—
a. Lanius collurio. b. Paradisea raggiana. C. Gymnorhina organicum.
Letters as in text-figs. 101, 102.
364 MR. W. P. PYCRAFT—CONTRIBUTIONS [Apr. 9,
stand out in sharp contrast with the ventral median keel of the
body. The horns rise directly upwards for a considerable distance,
and being continued backwards along each side of the dorsal
aspect, this region is converted into a deep triangle.
It is to be noted that in the matter of the vomer and its
relation to the maxillo-palatines, the skull of Hwryceros very
nearly approaches the Gymnorhine. This is a fact which greatly
assists in the difficult task of deciding the true position of this bird
in the system. It is further noteworthy that the peculiarly
inflated condition of the maxillo-palatines in the pre-maxillary
region may well have been derived from conditions such as obtain
in the Gymnorhine.
The pterygoid throughout this series is elongated, and round or
triangular in section. Basipterygoid processes being absent, this
rod extends directly from the quadrate to the palatine, and
parasphenoidal rostrum. Consequently the principal feature of
interest for description here concerns the nature of the palato-
pterygoid articulation. Before entering on this subject, however,
it may be remarked that the triangular flange which, in the higher
Passeres, e. g. Corvide, extends from the rim of the dorsal
segment of the cotylus for the quadrate, backwards along the
shaft, is in the forms herein described represented by a slender
style, forming a V-shaped angle with the shaft of the quadrate,
e.g. Sayornis. In many cases, however, it is reduced to a mere
tubercle, or it may even be absent.
As touching the nature of the pterygo-palatine articulation.
This takes two forms. Generally, the pterygoid terminates
distally in a leaf-shaped plate, the mner surface of whicii is
applied to the parasphenoidal rostrum, while the inferior edge
thereof glides over the edge of the obliquely truncated proximal
end of the palatine. This method of ar ticulation has been derived,
as I have elsewhere pointed out (6), from an earlier condition
of things wherein the pterygoid extended forward to support the
vomer. By segmentation of the pterygoid, and the fusion of the
vomerine portion with the palatine, the characteristic palato-
pterygoid joint has been formed. Ina number of cases, however,
a pseudo-primitive condition obtains: the pterygoid, instead of
segmenting, preserves its integrity, and fuses with the palatine,
e. g. Dicruride, Prionopide, Laniine, Gymnorhine, Malaconotide,
Artamidee, and Vangide. This fusion of the pterygoid and
palatine I have already shown (5) obtains also in the Hurylemide
and the Tracheophone Passeres.
The palatine, distad, has the form of a straight rod which is
continued backwards considerably beyond the level of the
antorbital plate, terminating in a more or less sharp point.
Immediately below the inferior border of the antorbital plate it
sends inwards, almost at right angles, a broad bar which gives rise
to a long trough-shaped scroll running parallel with the long axis
of the skuli, and with its concavity turned towards the middle line.
The upper edge of the scroll affords support to the vomer and
pterygoid, while the inferior margin is free, and with its fellow of
1907. | - TO THE OSTEOLOGY OF BIRDs. 365
the opposite side forms a long tubular cavity, open below, and
roofed above for the most part by the vomer.
In my earlier papers on the Eurylemide and Tracheophone
Passeres (5) I have described the upper lamina of this scroll as
affording a surface to facilitate gliding along the parasphenoidal
rostrum. As a matter of fact, this is not really the case: the
palatine hardly touches the rostrum, which is embraced only by
the distally expanded ends of the pterygoid.
The free end of the palatine rod—caudad of the bar which
connects this with the body of the bone—is conspicuously long
and spike-like in many Tyrannide, the Vangide, and Artamide
(if these two can be regarded as distinct families) and Gymnorhine.
This spur is referred by W. K. Parker to the “ transpalatine,” but
it is doubtful whether this homology exists.
The guadrate presents no very striking characteristics. It
differs from that of the HEurylemide and the Tracheophone
Passeres in that, except in the Phytotomide, the quadrato-jugal
bar is not set out from the quadrate by a cylindrical boss of bone
laterad of the outer articular condyle. The condyle, however,
stands far out, laterad, of the long axis cf the quadrate. The
internal mandibular condyle of the quadrate is, in the Phyto-
tomide, conspicuous for its large size and spherical shape. The
squamosal and otic heads are well marked ; and the orbital process
is conspicuous for its great length, terminating in a point, except
in the Phytotomide wherein thé orbital process is greatly reduced.
The Mandible.
The mandible, like the quadrate, affords no very important
features. The lateral vacuity, which is placed far back, is always
small. The angular process is always short, especially in the
Gymnorhine and Vangide, and in Pitangus among the Tyrannide.
In Artamus and Gymnorhina the mandible is truncated posteriorly.
The internal angular process is of moderate length, though in
Pityriasis and Artamia it must be described as very short.
The Hyoid.
Unfortunately in only three skeletons of the whole series
examined during the preparation of this paper are the hyoid bones
preserved. At the time when these skeletons were made no
great care was taken to preserve these bones. As these hyoids
belong to very different forms and have not hitherto apparently
been recorded, we may briefly indicate their structure.
The Dicruride are represented in this matter by Hdolius
Jforficatus. Herein basi-hyals are represented by a pair of rod-
shaped bones lying side by side, and terminating, posteriorly, in a
free spine. Basi-branchials 1-2 are fused, and both agree in
being much elongated. The cerato-branchials are of moderate
length ; the epi-branchials are unfortunately missing.
In Newtonia the basi-hyals are relatively longer, narrower, and
enclose a space between them. Basi-branchial 1 is shorter than 2,
Proc. Zoot. Soc.—1907, No. XXV. 25
366 MR. W. P, PYCRAFT—CONTRIBUTIONS [ Apr. 9,
which is relatively somewhat longer than in Hdoliws. The cerato-
and epi-branchials are of moderate length.
In Myiarchus the basi-hyal is wanting ; cerato-hyal 1 is short
while 2 is relatively longer than in the species just described.
The cerato-branchials are extremely long; the epi-branchials
unfortunately are missing.
i. THe VERTEBRAL COLUMN.
The presynsacral vertebre are all heterocelous and free.
As to the general characters of these vertebre there is but little
that can be said to any purpose. The odontoid ligament is
ossified throughout ; hypapophyses occur on the 2—4 cervicals, aud
extend from the eleventh cervical to the first thoracic.
The composition of the column, however, furnishes one or two
interesting points.
First, as to the cervical and thoracic regions. The last thoracic
vertebra has passed into the synsacrum, but, ignoring this fact
for the present, we may remark that the total number of vertebre
in these two series ranges from 19 to 21,
The last cervical bears a pair of long free-ribs, but without
uncinate, or sternal segments. As a rule there are but six
thoracic vertebre bearing long ribs, and of these vertebrae the
last is fused with the synsacrum, while its ribs just fail to reach
the sternum. Sayornis among the Tyrannide, however, has
seven thoracics. The sternal segments of the 6th pair are attached
to the sternal segment 5, some distance above the sternum, while
the similar segments of the 7th pair are attached to small facets,
one on each rib, high up on the sternal facet of the 6th. Of
the seven thoracics in Artamia the ribs of the 6th, pair reach
the sternum, articulating therewith immediately behind the
5th pair by a much reduced joint. The ribs of the 7th pair of
vertebre are represented only by a pair of rods, almost hair-like
in delicacy, attached to the tip of the thoracic, and the upper third
of the sternal segment of the 6th ribs.
The number of synsacral vertebre ranges between 16 and 19;
the number of free caudals between 6 and 8. In no case have
both the sacrals preserved ribs, and only rarely can traces of the
ribs of the 2nd sacral be found. The chief numerical differences
may be stated as follows :—
TYRANNID2. MUSCICAPID®.
Sayornis. Tyrannus. | Terpsiphone.
Thoracic ......... li 1 | a
Brumibar (02022. 2 3 >
Lumbo-sacral ... 1 2 1
Sacral yee ee 2 2, | 2
Caudalie 3+7 free caudal A+7 | 4+7
9 12 | 11
Total 16. Total 19s" “) ~ otal 18:
1907.) 10 THE OSTEOLOGY OF BIRDS. 367
DICRURID#. | GYMNORHINE.
Hdolius. | Pityriasis.
iVoracica eee eee 1 i
Barapa. pee ee: 3 3
Lumbo-sacral ............ il 2
POP CIT?T ARAM S aR aed tines MEER 2 2
Canadian. Bk ek eed 4 +6 | 5+6
11 | 13
Total 17. | Total 19.
Compared with the Tracheophone Passeres these numbers are
of some interest, inasmuch as they show that the vertebral column
of the forms now in question has become more specialised by the
reduction in the number of the synsacral and free caudal vertebre.
In the Tracheophonz the number of these vertebre is never less
than 19, the Jumbo-sacrals are never less than two—generally three
are present: as against the single vertebra of many Tyrannide,
Muscicapidee, and Dicruridee.
iv. THe Riss.
The number of the ribs has already been referred to, but a
brief reference may be made here to one or two structural pecu-
liarities of interest. In Pityriasis, for example, the upper portions
of the thoracic segments are conspicuous for their great breadth,
as well as for the unusually large size and breadth of the un-
cinates which are attached to their respective ribs by broad
oblong bases. — Buchanga, one of the Dicrurid, shows an approach
to this condition. Only in Gymmnorhina, apparently, has the
last thoracic rib retained its uncinate. It would be interesting to
discover whether there is anything in the mode of life of the
Gymnorhine which could account for the width of the thoracic
ribs and uncinates just referred to.
v. THe STERNUM AND SHOULDER-GIRDLE.
With the exception of the Pittidee the sternum is of the typical
Passerine type in all the forms herein described, having but a
pair of posterior notches and a long Y-shaped spina eaterna. The
peculiar sternum of Pitta has already been described by me (5).
Except that the corpus sterni varies in its relative length, the
sterna now under consideration afford no appreciable distinguish-
ing characters. Only the Gymnorhine and the Paradiseidz have
the sternal plate elongated, and even here this is nota very striking
feature.
The coracoid grooves in all cases are deep; and similarly, in
all cases, the articular surfaces for the sternal ribs are confined to
the anterior lateral processes.
The carina sterni is deep in all save the Gymnorhine, though
One
368 MR. W. P. PYCRAFT—CONTRIBUTIONS [Apr. 9,
even there it is fairly well developed, except in Pityriasis and
Huryceros, which have a markedly shallow keel.
The posterior notches, though differing slightly among different
genera, in the matter of length, do not afford any matter for
comment except in the Paradiseide, where, by the expansion of
the free end of the posterior lateral process, the notch becomes
converted into a foramen. Whether, however, this character
obtains throughout the Paradiseide remains to be seen.
The coracord shaft is typically passerine, being long and slender,
and having a broad flange extending along the outer border of
the shaft from the base forwards nearly as far as the middle ;
while the acrocoracoid is long, turns forwards and inwards, and
sends backwards a hook-shaped process affording additional attach-
ment for the furcula. The procoracoid is obsolete or absent,
except in the Tyrannide and Phytotomide, where it forms a long,
band-hke spur affording a shelf for the articulation of the
scapula.
The scapula aftords no matter for special comment except in
so far as the acromion is concerned. In the Tyrannide and
Phytotomidee this process is not so well developed as in the higher
forms herewith associated, masmuch as it does not send down,
into the foramen triosseum, between the clavicle and the coracoid,
a spur for the support of the free end of the furcula. Instead the
posterior border of the pedate expansion of the free end of the
furcula rides in a trough hollowed out of the anterior face of
the acromion. Inthe remaining forms here described this method
of articulation also obtains, but the inner wall of the trough is
continued downwards to form the spur to which allusion has
just been made.
The furcula presents a remarkably uniform character through-
out this series, varying, to any appreciable extent, only in the
form of the expanded free end of each limb. In all, the hypo-
cleideum is more or less oblong and directed upwards, so as to
present a nearly vertical face towards the anterior border of the
carina sterni. In the Tyrannidee and Hirundinide each limb
terminates in a roughly spatulate free end the postero-superior
angle of which is received into a groove in the acromion of the
scapula. The middle region of the outer surface of this plate
is applied to the acrocoracoid, thus enclosing the foramen tri-
osseum. By way of contrast a comparison should be made with
say the furcula of Gymnorhina. Here the free end is roughly
T-shaped, while it articulates with the anterior face of the acro-
mion rather than with its end, a relationship which, it may be
noted, may be seen in its incipient stages in the Tracheophone.
The foramen triosseum, then, in the Tyrannide is bounded,
without by the acrocoracoid, and on the inside by the expanded
free end of the furcula and the acromion of the scapula—these
two elements articulating by means of two opposed surfaces set
transversely to the long axis of the scapula. The higher Passeres
differ only in that the articulation between scapula and furcula
1907. ] TO THE OSTEOLOGY OF BIRDS. 369
is by oblique facets, so that the posterior half of the inner wall of
the foramen is bounded by a spine-like projection fashioned out
of the acromion,—the inner side of the spine forming the articular
surface for the furcula.
vi. THe Petvic GIRDLE.
The pelvic girdle presents no very striking modifications,
neither does it afford any important structural characters such as
can be of use for systematic purposes. But there are nevertheless
features which deserve mention.
In the Tyrannidz the pelvis is broad, the ilia being widely
separated by the transverse processes of the synsacrum. The
pre-iliac region is moderately long and varies slightly in its shape
in different genera. The superior margin of the fovea iliaca
anterior in all, however, fades away in the region of the anti-
trochanter and is not continued backwards into the free edge of
the dorsal plane. The postacetabular ila are broad, and twisted
horizontally to form a broad dorsal plane, and enclose between
them a broad plate formed by the synsacrum and its transverse
processes. Ossification however has almost completely obliterated
the intertransverse sacral foramina. The ischium is narrow, and
has its postero-inferior angle produced backwards into a long style,
which, at its tip, serves as a point of attachment to the pubis.
This is long and slender, and is produced backwards for some
distance beyond the free end of the ischium. The ischio-pubic
fissure is wide and long; shut off anteriorly from the obturator
foramen by a bony bar, and closed posteriorly by the junction
of the ischium with the pubis. The ilio-ischiadic foramen is
relatively large. There is no pectineal process. The fovea luwm-
balis is shallow, and the fovew ischiadica and pudendalis. pass
almost insensibly into one another.
The pelvis of the smaller forms of the Families now under
consideration does not differ materially from that just described
as typical of the Tyrannide. In larger birds, such as the Gym-
norhine, the girdle is relatively much longer, while the pre-ilia
rise up to the level of the synsacral ridge, nearly meeting one
another in the middle line. There is a single row of intertrans-
verse sacral foramina; the ischio-pubic fissure is very wide, and
closed posteriorly, but itis not shut off from the obturator foramen
by a bony bar.
That the form of the pelvis may be very largely modified by the
decline of the power of flight and the consequent increased use of the
hind limbs, may be seen in the pelves of such birds as Huryceros
prevostir and Newtonia brunneicauda. Both these species would
seem to be relatively poor fliers, inasmuch as the carina sterni is but
feebly developed: and this is especially true of Newtonia brunnei-
cauda—one of the Muscicapidee wherein the keel of the sternum
is remarkably shallow. Here the width of the pelvis is increased
by the lateral extension of its dorsal plane, while greater depth is
370 MR. W. P. PYCRAFE—CONTRIBUTIONS [ Apr. 9,
gained by the sharp downward turning of the spine-like postero-
inferior angle of the ischium which transforms the closed ischio-
pubic fissure into a foramen of considerable width.
vil. THe Pecrorat Lime.
Were it possible to examine the wing-skeletons of a large
number of genera of any of the Families herein described, data
might be found for use in systematic work. So far, however,
such a collection is not to be met with in any museum. From
the material at my present disposal it is possible to deal only with
the main features of the wing, and these do not display any very
remarkable characters.
The humerus only is pneumatic, except in the Hirundinide,
wherein all the wing-bones are non-pneumatic. But this
segment of the limb differs, in the several genera, chiefly in
the relative development of the deltoid crest, and of the ect-
epicondylar process: a relation, in so far as the deltoid crest is
concerned, which varies as the power of flight. Where this is
strong, as in the Hirundinide and Gymnorhine for example,
this crest is of considerable size, having an oval contour, pro-
jecting well beyond, and extending down the shaft, for about
one-third of its length. Where the flight is weak, as in the
Tyrannide, this crest is barely developed. Curiously, however,
the size of the ectepicondylar process does not appear to depend
on the flight to the same extent, since in 7yrannus, for mstance,
it 1s relatively larger than in G2 ymnorhind. But this appearance
is deceptive, and is due to the relatively more slender shaft of
Tyrannus. Nevertheless the ectepicondylar process does not
appear to be so directly under the influence of flight as the deltoid
crest. In Vylas, for instance (in the ‘ Hand- list of Birds’ placed
with the Seta but probably one of the Prionopide, or
Dicruride), the power of flight, judging from the size of the
carina sterni, does not seem to have undergone any serious re-"
duction; this process is small, and surmounted by a small, but
sharp spike. Similarly, in the Crmmpopbae te this process is
ereatly reduced, but a small spike remaining; and here again
full powers of flight are certainly retained. Tn this reduced « size
of the ectepicondylar process, these birds resemble the Pittidee
and Eurylemide, so that yet another alternative may be pre-
sented—the character may be a primitive one. In the Swallows
there is a large entepicondylar process lying behind the ulnar
condyle, and projecting directly backwards.
The forearm is relatively long and straight in all, though the
radius is slightly broad save for its distal one-third which is
straight. The ulna has prominent tubercles for the attachment
of the secondary remiges, and a pointed olecranon process, which
interlocks with a corresponding entepicondylar spur on the
humerus.
The manus is the most variable segment of the limb, inasmuch
1907. ] TO THE OSTEOLOGY OF BIRDS. 371
as in some forms, as in the Tyrannide for example, and in Vew-
tonia among the Muscicapidee, it does not exceed the humerus in
length, while in others, as in the Gymnorhine, it considerably
exceeds the humerus, being as long as, or slightly longer than,
the forearm. Pityriasis, however, among the Gymnorhine is
exceptional : all the segments of the wing being short, the fore-
arm especially so; but this, it is to be noted, is associated with a
relatively shallow sternal keel. The intermetacarpal plate is
well-developed, and as a rule there is a deep groove along the
dorsal aspect of Me. II. for the tendon of the extensor indicis
longius. This groove is exceptionally deep in Gymnorhina and
the Hirundinide.
In the Swallows indeed the manus has become considerably
modified in response to the great strains put upon it. And this is
especially noticeable with regard to Ph. 1 of D. II. This phalanx
is of great breadth, flattened dorsally, and widest at its distal end :
while on its ventral aspect it presents the appearance of an oblong
bone deeply hollowed along its centre, and having its preaxial
border produced proximad in the form of a tubercle to afford extra
ligamentary attachment to its metacarpal. Metacarpal III. is
produced for a considerable distance beyond Me. II. and supports
a slender phalanx. The terminal phalanx of D. II. (Ph. 2) is
very small. In the Hirundinide the humerus does not exceed
the carpo-metacarpus in length. The pollex is greatly reduced.
Compared with the Kurylemidz the wing, in the forms herein
described, shows a more specialised condition in the greater devel-
opment of the deltoid crest and ectepicondylar process of the
humerus ; and in the broadening of the metacarpals I. & II.
vill. Tae Petyrc Limes.
When the skeletons of a sufficiently large number of genera
and species have been got together, it will probably be found that
the pelvic limb will reveal useful characters for systematic pur-
poses. With the limited material at my disposal however, I can
do no more than outline the general characters of this portion of
the skeleton.
The femur, which is relatively long and slender, appears to be
pneumatic only in Zylas, Newtonia, Huryceros, Artamia, and
Artamus. The popliteal fossa is obsolete. The tibial condyle
develops a more or less prominent spur laterad of the slight de-
pression which marks the popliteal depression in Vewtonia, Vireo-
lanius, and Paradisea. Asa rule the tibial and fibular condyles
are subequal, but in the Gymnorhine and the Dicruride the tibial
condyle is markedly larger.
The tibio-tarsus is long and slender, and slightly inflected at
the tip: the intercondylar gorge is deep and wide; while the
extensor groove immediately above is traversed by a bony “ ex-
tensor bridge,” which is generally wide. Immediately above the
bridge, on the fibular side of the shaft, is a more or less prominent.
372 MR. W. P, PYCRAFT—CONTRIBUTIONS [ Apr. 9,
tubercle, while on the opposite side of the tibial shaft, and some
considerable distance above the entocondyle, is a similar condyle,
generally much larger. These serve as points of attachment for
the ligamentum transversum, which keeps in place the tendon
of the tihialis anticus. The ento- and ectocnemial crests differ
considerably in their relative development. Thus in the Tyran-
nide they are but feebly developed ; in the Pittide the ento-
cnemial is large, the ectocnemial very small; while in the
Gymnorhine, for example, both are very str ongly developed.
The size of these crests does not appear to vary in proportion as
the wings are less used. Buta more careful study of the habits
of the different types which present these differences may show
that they vary as the legs are used in perching—as in birds
haunting thick undergrowth and flying but little—or walking.
The tarso-metatarsus is short in arboreal types, such as the
Tyrannide, long in those which spend their lives for the most
part on the ground, as in the Pittide and Gymnorhine. In all,
the outer border of ‘the shaft is produced backwards into a bladder-
like ridge; and in some this is very marked, as in Gymnorhina
and Paradisea for instance. The hypotarsus is complex, and
takes the form of a backwardly projecting spur pierced by several
foramina.
The trochlee are, as it were, cut out of the flattened and
expanded distal end of the shaft, so that all lie in the same plane.
The ento- and mesotrochlea are subequal in length, but the ecto-
trochlea is slightly shorter, and generally also much smaller: pre-
senting only a narrow rim for the articulation of the digit.
Only the mesotrochlea indeed presents the typical grooved
articular surface.
The anterior face of the proximal end presents characters which
may prove of service. Thus in the Tyrannide, for instance, this
region is rounded on its outer border, and bears a tubercle for the
insertion of the tibialis anticus ; while below the inner cotylus is
a bony bridge—the ossified ligament which guides the tendon of
the eatensor longus digitorum. In the Gymnorhine this region
of the shaft is deeply hollowed: the extensor bridge is ligamentous,
and the shaft is perforated by two small foramina, one on either
side of the median digit. The Pittide appear to agree with the
Tyrannide in these particulars, which appear to obtain also
throughout the rest of the Families herein described.
1x. SUMMARY.
I have ventured, on the evidence chiefly of osteological and
myological characters, to associate many Families which have
hitherto been separated, often widely. Furthermore I have en-
deavoured, in the course of this summary, to show that the ranks
of these Families must be enlarged to receive forms which till now
have been regarded as having no affinities therewith.
It will be remarked that I have included im this paper the
1907. | TO THE OSTEOLOGY OF BIRDS. SS
Families Phytotomide, Pittide, and Tyrannide. This I have
done because there seems good reason to regard these three
nearly related Families as close allies of the Diacromyodean
Passeres.
As Fiirbringer—improving on the earlier work of Huxley (2),
Sclater (7), and Newton (3)—has already suggested, the Passeri-
formes should, I think, be regarded as forming: four Suibarders 2 —
1 Eurylemi.
2 Oligomyodi.
3 Tracheophones.
4 Diacromyodi= Oscines.
With the Hurylemi must be included the Hurylemide,
Cotingide, Pipridz, and Philepittide: the whole forming one of
four great branches of a common stem. The other three give
rise, respectively, to the 'Tracheophone, the Tyrannine forms
(including the Phytotomide and Pittide), and the Diacromyodean
Passeres.
The Phytotomide, I would remark, are certainly more nearly
related to the Tyrannide than to the Pittide; and this fact is
largely borne out by the structure of the skull.
Osteologically, the Oigomyodean and Diacromyodean suborders
have much in common. The skeletons of the Tyrannide and of
the Muscicapidze, for instance, present very close resemblances.
The formation of the Suborders just referred to is based largely
on myological characters, such as are afforded by the syrinx and
the wing-muscles. All that concerns the former is so well known
that it will not be necessary to repeat it here. As to the wing-
muscles, however, a few words are necessary.
In earlier papers (5) I have already shown that the deltoideus
major (pars longa and pars brevis) affords useful characters for
systematic purposes; and the condition of these muscles in the
groups now under discussion offers further confirmatory evidence.
Thus, put briefly, in all save the Diacromyodean Passeres the
pars brevis fails to reach the ectepicondyloid process of the humerus.
In the Eurylemid group it is very short, being confined to the
upper end of the humerus; and this is true also of the Pittide
in the Oligomyodean evoup : ; but in the Tyrannide this muscle
extends more than halfway down the shaft. In this the Tyran-
nide agree with the Tracheophone Passeres, from which however
they differ in the syringeal muscles, osteology, and pterylosis.
Nevertheless, the differences between the wing-muscles in the
Tyrannide and Pittide are differences of degree and not of kind.
Diacromyodean Passeres alone seem to have retained the del-
toideus brevis in its entirety: but even here it shows, in some
Families, a very marked tendency to decrease, while, in others,
on the other hand, it is relatively of great size. In the Musci-
capidee and Hirundinide it would seem to have preserved its
most primitive character. In the former, this muscle is very
thick and fleshy throughout its whole length up to the point of
374 MR. W. P. PYCRAFT—CONTRIBUTIONS [ Apr. 9,
insertion, where it becomes suddenly tendinous. In the Hirun-
dinide it is also of great size, and joins the deltoideus major
longus to form a short, thick, fleshy column to be inserted into
the ectepicondylar process. The d. m. longus is a relatively
slender muscle in both these families; and this is true also of
the Gymnorhide and Paradiseidee.
The Families brought together in this paper, are, it is con-
tended, all more or less closely allied. And this relationship is
nowhere more easily traced than in the skull.
Throughout this series, the maxillo-palatme processes retain
more primitive characters than in any of the remaining Diacro-
myodean Passeres : and these would seem to have been derived from
the Pittide and Tyrannide. In the Tyrannine palate, as has
already been remarked, the maxillo-palatines take the form of a
pair of broad triangular plates; the apex of the triangle tending
to curve towards the base of the skull.
The Diacromyodean types show an ever increasing tendency to
transform this triangular plate into a long slender rod, curving
towards the base of the skull, and terminating in a more or less
spatulate, and sometimes inflated free end.
The Campophagide and Muscicapide well illustrate these
phases, the more primitive showing the triangular, the more
specialised the rod-like form, Other Familes herein described
show further and more peculiar characters of the palate which
will be more particularly referred to in due course.
Broadly speaking, the Families now under consideration may be
divided into two Groups—F lycatchers and Shrikes.
The Flycatchers (Muscicapide and Campophagidee) most nearly
approach the Tyrannine type; and are therefore to be regarded
as the more primitive. Herein the maxillo-palatines are more or
less triangular, and there is an ossified septum nasi dividing the
palatine foramen into right and left moieties. Commonly, the
inferior border of this nasal septem is expanded to form a
flange.
The Hirundinidz are generally regarded as very nearly allied
to the Muscicapidee. That they are related to this Family is,
I think, a fact; but this alliance does not appear to be so
close as is generally supposed. The Swallows indeed are a race
apart.
In other words, the Swallows and Flycatchers are to be
regarded as derivatives of the same generalised stock which have
developed along independent, and more or less parallel lines.
Tn so far as the skull is concerned, the Swallows recall some of the,
presumably, more primitive Tyrannide. And this is especially
the case with regard to the specialised condition of the maxillo-
palatines and the relation of the palatines to the premavxilla:
characters which appear to be, to some extent, adaptive, and
correlated with the shortening of the beak and widening of the
gape which apparently becomes necessary to ensure the successful
1907.] LO THE OSTEOLOGY OF BIRDS. 375
capture of small prey when on the wing. In the matter of the
deltoideus muscles the Swallows do not differ greatly from
the Muscicapine forms, but in their pterylosis they differ from
all other Passeres, in that the pteryla spinalis is forked. This
condition, however, may well have been derived from the breaking
up of the lower segments of an elliptical tract enclosing a space,
such as is seen in the Tyrannide, or the Muscicapide for
example.
We may pass now to what I have regarded as the Laniine
forms. These seem to be divisible into two groups including,
among others, the following families of Dr. Sharpe’s Hand-list:—
Laniide, Prionopidee, Artamidz, Vangide, and Alrocharide. The
Dicruride must also be included here, for they show, on the one
hand, affinities with the Malaconotide, and on the other with the
Prionopide. But many of these Families, as at present constituted,
contain genera that cannot be allowed to remain there. While
genera now included in other Families, outside this Group, will
probably have to be transferred thereto.
We shall probably be near the truth in regarding the Laniidee,
Prionopide, and Malaconotide as representing so many branches
of acommon stem. Of this stem the Gymnorhine group must
apparently be regarded as a sister branch, similarly splitting
up into three branches—the Gymnorhine, Artamine, and
Vangiine (if these two can really be separated), the Vireolanide,
and probably the Paradiseidee.
This Gymnorhine group may be distinguished by many
characters, the most striking of which are afforded by the skull—
the peculiar form of the palate, and the long postorbital processes
already described in the earlier pages of this paper.
From this Gymnorhine branch it is probable the Paradiseide
are derived. This latter Family, by the way, as it now stands,
contains many forms which must be placed elsewhere. The
Bower Birds (Ptilonorhynchide), for example, do not seem to
possess any claim to be included here.
The distinctness of the Gymnorhine group has been ignored by
many later workers, but there seems good reason to believe that
W. K. Parker was justified in proposing therefore the term
“* Austro-coraces.”
As touching the Vireonide. This Family is regarded apparently,
by some as allied to the Sylviude, by others to the Laniidee.
The latter view is certainly the more correct. But so far as my
material goes, everything points to a near relationship to the
Muscicapidee. Of the six genera, however, usually included in
this Family two at least must be removed; for Cyclorhis would
seem to be very nearly allied to the Laniide, while Vireolanius is
almost certainly closely related to the Artamide. This genus is
probably an early offshoot of the branch which eventually gave
rise to the Artamide, and may therefore either be included in
this Family, or be allowed to rank as a separate Family; better
316 MR. W. P. PYCRAFT—CONTRIBUTIONS - [ Apr. 9,
still perhaps the Artamide should be split up into two sub-
families—Artamine, Vireolaniine.
I have been much puzzled as to the systematic position of the
genus Sphecotheres. Generally regarded as one of the Oriolide, it
seems to me much more nearly allied to the Campophagide, and
should indeed be included in this Family. The skull bears a
general resemblance to that of Graucalus. It is certainly not an
Oriole; at any rate if the skulls in the Museum Collection are
rightly labelled, and there seems to be no reason to doubt this.
The fact that Sphecotheres has not developed the peculiar spiny
rump-feathers so characteristic of the Campophagide may be
urged, by some, as an objection to the introduction of this genus
to the Family Campophagide. This, however, does not seem a
very weighty objection, for the feathers in question vary in the
degree of their spininess very considerably, in some genera it 1s
hardly noticeable.
The genus Zylas again—which includes five species—has
puzzled me greatly. Included by Dr. Sharpe with the Pyeno-
notide, it seems to me to belong rather to the Prionopide, though
in some respects it recalls the Muscicapide. It has certainly no
affinities with the Pyenonotide.
The resemblances which the birds of the genus TZylas
undoubtedly present, in many osteological characters, to the
Laniine group are significant ; and this because, curiously enough,
they have been, and still are, regarded as “ Babblers” (Pycno-
notidz) which have, for purposes of mimicry, assumed the guise
of Shrikes !
Briefly, the genus Zylas includes five species, all of which are
peculiar to Madagascar. All bear a strong likeness one to the
other; and resemble scarcely less closely a member of the genus
Xenopirostris—X. pollent. This bird is considered by some to be
a Shrike, while by others it is, and probably rightly, included
among the Artamide, or the closely allied Vangidee—the two
Families must I think be united. There are three species in the
genus Yenopirostris, two of which are dull-coloured and bear no
likeness to _Y. pollent. Thus all the five species of 7’ylas seem to
have more or less completely assumed the livery of XY. pollent.
On the assumption that the genus Zylas belonged to the
Pyenonotide, this resemblance to an agressive Shrike was truly
remarkable; but the likeness loses little of its interest even now
that it is shown that this genus really belongs to Prionopide, a
Family which must be included among the Shrikes.
Finally, I propose to arrange the Diacromyodean Families
herein described in four groups, as follow :—
Suborder D1acrRomyopt.
Group I. Hirundines.
Fam. 1. Hirundinide.
1907.] TO THE OSPTEOLOGY OF BIRDS. 377
Group IT. Muscicape.
Fam. 1. Muscicapidee.
2. Campophagidee.
3. Vireonide (excluding Cyclorhis
and Vireolanius).
2?
Sz)
Group IIT. Laniv.
Fam. 1. Lanude.
2. Malaconotide.
3. Prionopide.
4. Dicruride.
5. Cyclorhide.
Group IV. Gymnorhines.
Fam. Gymnorhide.
Artamidee.
Vireolanide.
Abvocharide.
. Paradiseide.
2?
99
7?
On = (SU) i) m
29
Text-fig. 104.
Syrr PRLPUAR.
\ Wialacoroe ae
Arlarnide. Prtonoprae.
Deerurrde| Lar x,
VALZAX cretefat ae
Utreolaruiiude
A; ochartide. Cyclo: Harundirntar
Vrreo7t Le
Pecradiserde.
Tyrac.
Phyloloyntde-
Pithde.
V4 ka Wlentrtaee.
Phylogenetic tree indicating the probable relationships of the Passeres Oligomyodii
and Diacromyodii.
378 CONTRIBUTIONS TO THE CSTEOLOGY OF BIRDS. [Atos 95
x. Key.
The following “ Key” gives the cranial characters of the
Families dealt with in this paper.
Suborder OLIGOMYODI.
Maxillo-palatines triangular; vomer truncated; nasal septum partly ossified ;
post-orbital process reduced; processus zygomaticus squamosi large; quadrate
with a lateral pillar for articulation of the quadrato-jugal bar; a shallow
temporal fossa extending as far back as the middle of the lambdoidal ridge.
Tyrannvide.
Maxillo-palatines in the form of broad linguiform plates sending up a slender spur
into the lachrymo-nasal fossa; anterior palatal foramen nearly closed; maxilla
with tubercles along the tomium; post-orbital process obsolete; processus
zygomaticus squamosi greatly developed; quadrate with a vertical, lateral pillar
for articulation of quadrato-jugal bar ............................... Phytotomide.
Maxillo-palatines in the form of laminate spurs; vomer large, and deeply cleft
posteriorly ; post-orbital process small; processus zygomaticus squamosi large ;
nasal septum ossified: beak with a nasal hinge; temporal fosse separated by a
moderately broad sagittal crest ................:eeec ete, Pittide.
Suborder DIACROMYODI.
Group J. HirUnDINEs.
Maxilla passing abruptly into the quadrato-jugal bar in the region of the
antorbital plates, projecting laterad of the bar in the form of a wide ledge.
Maxillo-palatines slender rods with spatulate free ends underlying the
vomer. Nasal fossa large, septum nasi wanting; anterior palatine foramen
of great size; external aperture of ear large, oval, directed downwards and
outwards. Post-orbital process feebly developed ...... Hirundinide.
Group II. Muscicars,
Mavillo-palatines produced into flattened bands. Post-orbital process feebly
developed, low down, and separated by a notch from the processus
zygomaticus squamosi which is feebly developed. Temporal fossa greatly
reduced or wanting.
a. Nasal septum ossified .....................0ee eee. DMuscicapide.
8. Nasal septum wanting
a’, Post-orbital process prominent; maxillo-palatines
with hastate free ends ..... ................0.04.00....... Campophagide.
8’. Post-orbital process obsolete; imavxillo-palatines
with spatulate free ends ........ .........-.............. Vireonide.
Group II], Lantt.
Maxillo-palatines in the form of short, more or less triangular spurs with a
palatal vacuity.
a, Post-orbital process feebly developed; processus
zygomaticus squamosi large; nasal septum strongly
developed: hcxictaccossccc to teecteece | LaCTLaees
(3. Post-orbital process well developed ; processus zygo-
mMaticus squamosi large; nasal septum imperfect ;
maxillo-palatines embracing facets on the vomer ... Malaconotide.
y. Post-orbital process well developed ; processus zygo-
maticus squamosi moderately large; nasal septum
well developed; maxillo-nalatine processes embracing
THAGIS ON WA WONG nosesecobccoccrs-eosetooosuccooacconecvann JDUCRIMPHU
6. Post-orbital process well developed ; processus zygo-
maticus squamosi obsolete; nasal septum nearly
TUN DTAES ORIEN AN WEXOWONINY cos sogescous cao ncasno coo san senncocses
¢. Post-orbital process obsolete; processus zygomaticus
squamosi large, conical; nasal septum feebly ossified ;
palatal vacuity nearly filled by septum nasi, and
longitudinal osseous bar forming floor of vestibulum. Cyclorhide.
Prionopide.
1907.] THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 379
Group IV. GymyNorwinus.
Palatal vacuity filled up by ossification of floor of vestibulum and nasal septum ;
maxillo-palatines short hastate spurs underlying vomer.
a. Post-orbital process and _ processus zy gomaticus
squamosi well developed, the Hommes low down and
separated from zygomatic process by a deep notch ;
mesial of zygomatic process of squamosal is a long
slender style; palatines with long spinous processes;
nasal fossa partly closed by ossification ; anterior
narial aperture reduced to a small oval................... Gymnorhide.
3. Post-orbital process large; zygomatic process of
squamosal small, or obsolete; the style mesiad
thereof feebly developed ; palatines with spinous
processes, short or obsolete; nasal fossa partly
closed ; narial aperture small, circular, lying in front
of a larger Ov! EXO — cooonnaspsonennana wsanaoanecanunaced AlpaaDOl@.
y. Post- orbital process obsolete ; ‘zygomatic process of
squamosal feebly developed; nasal fossa open......... Vireolanide.
6. Post-orbital process large ; zygomatic process of squa-
mosal small; nasal fossa closed; narial aperture
small, oval in shape; beak enormously inflated; a
wide foramen, ventrad and distad of miavillo-
palatine processes, leading into the air-chamber,
which gives the inflated appearance to the beak ...... Mrocharide.
¢. Post-orbital process obsolete; zygomatic process of
squamosal well developed ; palatines with very short
maxillo-palatal processes; nasal fossa small, but
co-extensive with narial aperture; with an imperfect
TORII LHD PEVDsb vinod ondoos coca oabodonosmenvedndobeomhentensdenedea . LeUMACIGEnelcas
REFERENCES.
(1) Fursrincer, M.—Untersuch. zur Morphol. u. Syst. des
Vogel. II. Allgem. Theil, 1888.
(2) Huxuey, T. H.—“ On the Classification of Birds.” P. Z.S8.
1867.
(3) Newton, A.—Art. “Ornithology.” Eney. Brit. vol. xviii.,
1884.
(4) Parker, W. K.—“ On Afgithognathous Birds.” ‘Tr. Z. 8.
Penang Ul woll, tea. Were
(5) Pycrarr, W. P.—“ Contributions to the Osteology of Birds.”
Part ViI., P. ZS. 19055 Part VIII, P.Z.S. 1906.
(6) Pycrarr, W. P.—‘‘Some points in the Morphology of the
Palate of the Neognathe.” Journ. Linn. Soc. vol. xxviii.
(7) Sctarer, P. L.—Cat. "Birds Brit, Mus., vel xiv., 1888.
(8) Saarve, R. B.—Hand-list of Birds, walls 1901,
April 23, 1907.
Dr. J. Rost Braprorp, F.R.S., Vice-President,
in the Chair.
The Secretary submitted the following report on the additions
that had been made to the Society’s Menagerie in March 1907 :—
The registered additions to the Society’s Menagerie during
the nian of March were 107 in number. Of hess 66 were
380 MR. R. LYDEKKER ON THE [ Apr. 23,
acquired by presentation and 3 by purchase, 28 were received
on deposit, 1 in exchange, and 9 were born in the Gardens. The
total number of departures during the same period, by death and
removals, was 163.
Amongst the additions special attention may be directed to :—
A pair of young Hippopotamuses (Hippopotamus amphibius)
from German East Africa, purchased on March 2nd.
An Eland (Zaurotragus oryx), born in the Menagerie on
March Ist.
A Banksian Cockatoo (Calyptorhynchus banksi) from New South
Wales, deposited on March 15th.
Dr. A. Smith Woodward, F.R.S., F.Z.S., exhibited an antler of
a Red Deer which had become malformed and enlarged by disease.
The specimen was obtained by Mr. Thomas Sheppard from a
prehistoric peat-deposit at Mablethorpe, Lincolnshire.
Mr. R. I. Pocock exhibited, on behalf of the Secretary, a model
of the African Elephant “‘ Jumbo,” formerly living in the Society’s
Menagerie, made by the late Mr. William Prehn and presented to
the Society by his widow.
My. C. J. With of Copenhagen communicated a paper entitled
“ An Account of the South-American Cheliferine in the Collections
of the British and Copenhagen Museums.”
This paper will be published entire in the ‘Transactions.’
The following papers were read :—
1. The Ears as a Race-Character in the African Elephant.
By R. LypEKKER.
[Received April 5, 1907. |
(Text-figures 105-121.)
That an animal with the immense geographical range of the
African Elephant—a range extending from Nubia and Abyssinia
in the north to the Cape of Good Hope in the south, and from
the east to the west coast across the heart of the continent—
should exhibit local differences is a practical certainty, even though
it be admitted that the animal is naturally a wanderer. Such
wanderings must, however, of necessity be limited in degree, and
are not of the ‘‘ Cape to Cairo” character which would be essen-
tial to cause uniformity in physical characteristics among all the
local forms of the species. The existence of local variation im
the species has, indeed, been well known for many years alike to
sportsmen, naturalists, and ivory-dealers; and in 1890 Dr. Paul
1907. ] EARS OF AFRICAN ELEPHANTS. 381
Matschie * proposed distinctive names for four of these local
races.
In the discrimination of these races Dr. Matschie relied chiefly
upon the characters afforded by the ears, supplemented by others
derived from the skull; and it is quite obvious that in the case of
an animal of the stature of 7 lephas africanus some such limitation
of the bases of comparison. Is a practical necessity, seeing that
entire mounted specimens are few and far between in our
museums, while even mounted heads are comparatively uncommon.
Unfortunately no figures of either the ears or the skulls have
been published, so that it is in some instances a matter of con-
siderable ditheulty to satisfactorily identify Dr. Matschie’s local
races.
Under these circumstances, | have thought it advisable to make
an attempt at putting matters on a more satisfactory footing by
comparing the ears of such examples of the African Elephant as
are available, either in the form of actual specimens or in the
shape of photographs, and to figure some of the more striking
types. I have taken the ear as the standard of comparison pri-
marily for the reason that it was specially selected by Dr. Matschie ;
and, secondly, because, with the exception of the tusks, it is the
one external portion of the animal which seems best suited for
this purpose, and is likewise one of which examples are not un-
frequently brought home by sportsmen.
The distinctive features afforded by the ear are exemplified not
only by variation in the matter of contour, but likewise by dif-
ferences in relative (and also in absolute) size. Why such
differences should exist (or why, for that matter, the ear of the
African Elephant should in all cases be so much larger than that
of its Asiatic cousin) it is hard indeed to divine; but that they do
exist, and ina very marked degree, will be apparent from the figures
in this paper, all but one of which are reproduced from photographs.
Then comes the question whether they are locally constant. To this
question I cannot give an absolutely decisive answer. The speci-
mens which I have had the opportunity of comparing agree,
however, respectively with Dr. Matschie’s descriptions of the
typical examples from the same localities; while the numerous
examples of the Sudan or Abyssinian Elephant which have come
under my notice, and likewise several of the Cape Elephants, all
conform in the matter of the size and shape of their ears to
their respective local types. So far, therefore, as the available
evidence goes, the various local representatives of the African
Elephant do seem to be fairly constant in this particular; so that
the onus of proving the opposite of this rests, I venture to think,
with those who may take exception to the views here advanced.
In the matter of the relative size of the ear, it is important to
* Sitz.-Ber. Ges. Naturfor. Berlin, 1900, pp. 189-190; a summary of the external
characters of these races is given by the Hon. Walter Rothschild in an appendix
to Major Powell-Cotton’s ‘A Sporting Trip through Abyssinia,’ London (Rowland
Ward, Ltd.) 1902.
Proc. Zoou. Soc,—1907, No, XX VI. 26
382 MR. R. LYDEKKER ON THE [ Apr. 23,
notice that this appears to be subject to a certain amount of
variation according to age; the proportionate size (as I infer from
the specimen now in the Society's Menagerie) being greater in
young animals than in those of riper age. It is thus evident that
for the purpose of defining local phases of the species, comparison.
should be restricted, so far as the matter of size is concerned, to
subadult or adult animals.
The following are the actual specimens that have come under
my notice :—Ist, the young Abyssinian Elephant now living
in the Society's Menagerie; 2nd, a mounted head from the
Lake Rudolf district, in the British Museum; 3rd, an entire
mounted specimen from Fort Manning, N.E. Rhodesia (South
Nyasaland), in the British Museum; 4th and 5th, two mounted
heads in the Imperial Institute, one from Mashonaland, and the
other reported to be from Swaziland; 6th, the right ear of an
Elephant shot in Congo Territory by Major Powell-Cotton, which
forms the fons et origo of the present communication. Mr. F. C.
Selous has lent me the ears of an Elephant shot by himself im
Mashonaland, one of which is now exhibited; and I am also
enabled to show, through the courtesy of the owner, the right
ear of a huge Elephant killed by Mr. A. Haig on the Blue Nile,
and the head of a male Somali Elephant belonging to 8.A.R.
le Due d’Orléans. The head of an Kast African Elephant with
very long tusks in Mr. Rothschild’s museum at Tring came under
my notice after the paper was read.
Of all these specimens, except the first and last, photographs
are herewith exhibited, and I may take this opportunity of
thanking the Director of the Imperial Institute for permission
to photograph the two heads under his charge.
In addition to the above, I have been provided with the follow-
ing photographs :—Ist, a South Somali Elephant in the act of
charging, photographed by Mr. R. McD. Hawker ; 2nd, “ Jumbo ”
and the ‘ Queen’s Elephant,” both formerly living in the
Society’s Menagerie; 3rd, an Elephant from the Aberdare Moun-
tains, in the private museum of Mr. 0. V. A. Peel at Oxford;
Ath, the photograph of the head of an Elephant from the Galla
Country, belonging to Sir EK. G. Loder; 5th, the head of an
Elephant shot by the Duke of Westminster in North-west
Rhodesia, now mounted at Eaton Hall; 6th, the head of an
Elephant shot by H.R.H. the late Duke of Saxe-Coburg-Gotha in
Cape Colony, now at White Lodge, Richmond ; 7th, the head of
a female South African Elephant in the Museum at Cape Town ;
8th, the head of a female of the same race in the Museum at
Saffron Walden; 9th, two Elephants from the Addo Bush, near
Grahamstown. For No. 3 I am indebted to Mr. Peel, for No. 4
to the Duke of Westminster, for No. 5 to Rowland Ward Ltd.,
for No. 6 to the Director of the South African Museum, for No. 7 to
the Curator of the Saffron Walden Museum, and for No. 8 to the
Director of the Grahamstown Museum; and to all these donors
my best thanks are due.
1907.] EARS OF AFRICAN ELEPHANTS. 383
After reading the paper, I received from Mr. Frederick Gillett
the photograph of an Elephant shot by himself in the Arusa Galla
Country (Long. 41° E., Lat. 7° 30' N.).
Before going farther, it may be well to mention that some
additional difficulty has been introduced into the work of com-
parison, owing to the ears of some of the specimens being in the
“cocked” and others in the recumbent position, and likewise
owing to certain differences in the orientation of the photographs.
To these difficulties, which I have endeavoured so far as possible
to discount, must be added any that may be due to vagaries on the
part of the taxidermists who have set up the various specimens.
Text-fig. 105.
Head of the Addo Bush, or Hast Cape Elephant (Hlephas africanus capensis),
from an adult male specimen in the Grahamstown Museum.
With these preliminary remarks, attention may be directed to
Dr. Matschie’s description of the ears and other external character's
of his four races. A free translation of the original paper enables
these to be given as follows, viz. :—
I. In the Southern race (Hlephas africanus capensis) the ears
are enormous, Somewhat square in shape, with rounded corners,
and a small, distinct, sharply pointed angular lappet in front. The
267
384 MR. R, LYDEKKER ON THE [Apr. 23,
forehead falls away towards the temples, so as to appear highly
arched.
Il. The Western race (1. africanus cyclotis), typically from
South Cameruns, also has the ears very large, but of quite different
shape, the contour being oval, and the lappet in the form of a
half-ellipse. The skin has a mosaic-like appearance, and its
colour is a paler grey than in the third race.
Tif. In the Sudan race (7. africanus owyotis) the ears are
considerably smaller and semicircular in shape, with the front
lappet very sharply pointed and angular.
IV. The East African race (4. africanus knochenhauert),
typically from German East Africa, has still smaller ears, which
are triangular in shape, with the front lappet angulated and
pointed.
In addition to these, a dwarf race of Klephant from the Congo
(Z. africanus pumilio), which may not have exceeded 7 feet in
height, has been named by Prof. T. Noack *; while the Albert
Nyanza Hlephant has been separated by myself as a distinct
race, under the name of #. africanus albertensis, characterised by
certain peculiarities in the form of the skull, which is unusually
short and broad.
Dr. Matschie, it should be added, was of opinion that the
Congo and the Angola Elephant might also be racially distinct,
while two or three other races might be represented in other parts
of Africa.
Here a word of explanation may be given with regard to the
element indicated by the word “ lappet” in the foregoing defini-
tion of Dr. Matschie’s races. This, I take it, refers to the
antero-internal angle of the ear, which forms a more or less dis-
tinct lobe, and, as in text-fig. 106, may be inflected.
As Dr. Matschie commenced with the South African EKlephant,
the same course may be followed on the present occasion. Here,
however, a difficulty at once presents itself, for there appear to
be two distinct southern forms of the species, one from the eastern
and the other from the western side of Cape Colony. It was to
the Hastern form, as represented by an Elephant from the Upper
Orange River district, that Cuvier gave the name of Hlephas
capensis; and since specimens from the same district afforded
Dr. Matschie the materials for his definition of the race described
under that name, we must apparently accept this determination
of the race, which may, however, really be inseparable from
Blumenbach’s /. africanus typicus, based on teeth from a locality
unknown.
A male Elephant from the preserves in the Addo Bush near
Port Elizabeth, mounted in the Grahamstown Museum, of which
the right side of the head is shown in text-fig. 105, agrees in the
squared form of the ear exactly with Dr. Matschie’s definition of
* Zool. Anz. Leipzig, vol. xxix. 1906, pp. 631-636.
+ “The Field,’ vol. cv. 1906, p. 1089.
1907.] EARS OF AFRICAN ELEPHANTS. 385
FE. africanus capensis, and may accordingly be referred to that
race. The inner front edge of the lappet is turned in towards
the neck. The high arching of the profile of the forehead
referred to by Dr. Matschie is strongly pronounced in the photo-
graph, as it is in the photograph of a second and younger specimen
in the Grahamstown Museum, which agrees precisely in the form
of the ears with the first example. I may add that the Director
of the Grahamstown Museum, to whom I am indebted for these
photographs, has been long convinced of the racial distinctness
ot the Addo Bush Hlephant, and has given me the following
additional particulars of the animal. The ears, compared to those
of the next race, are small, and the fore-legs proportionately low ;
while the ventral line of the body is almost. horizontal. More
important is the presence of a dense coat of hair on many parts of
the body ; this bemg very noticeable in the specimen of which the
skeleton is now mounted in the Oxford Museum.
SS AEN
Head of Female West Cape Elephant (#lephas africanus towotis), from a specimen
in the Sonth African Museum, Capetown.
Tf we call the preceding form the Addo Bush, or Hast Cape
Elephant, the race to which attention 1s now directed may be
termed the West Cape Elephant. That it is perfectly distinct
386 MR. R, LYDEKKER ON THE [ Apr. 23,
from FH. africanus capensis is, 1 think, certain, the ears being
relatively larger, and not having the slightest tendency to a squared
form. This is well exemplified by the photograph of the head of
an entire female specimen in the South African Museum at Cape-
town (text-fig. 106), which may be taken as the type of this race.
The ears are very long, somewhat in the form of a half-oval or
perhaps of half a pear, with the lappet moderately large, not
markedly pointed, but strongly inflected towards the neck. Appa-
rently the ears do not quite meet, when in repose, in the middle
line of the neck, and they are much larger in proportion to their
width than in the Addo Bush Elephant, in which, as already
mentioned, they approach a square. So far as I can learn, the
skin is not hairy.
“th
alike
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IM)
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Head of Male West Cape Elephant (Hlephas africanus towotis), trom a specimen shot
by H.R.H. the late Duke of Saxe-Coburg-Gotha in South Africa, and now at
White Lodge, Richmond.
Unfortunately, I have not the dimensions of the ears of either
the Cape Town or the Grahamstown specimen. Livingstone in
his ‘ Missionary Travels ’* has, however, recorded that in a female
elephant standing 8 feet 8 inches in height, the ears measured
4 feet 5 inches in vertical depth by 4 feet in horizontal diameter.
* Small edition, page 370.
1907. ] EARS OF AFRICAN ELEPHANTS. 387
While indicating the large size of the ears characteristic of South
African Elephants generally, these dimensions are suggestive of
the Addo Bush type.
Reverting to text-fig. 106, 1t may be noticed that the arching
of the forehead is less marked than in the Addo Bush Elephant.
Nearly similar features are displayed in the photograph of the
head of an immature South African Elephant in the Museum at
Saffron Walden, Essex, which was acquired somewhere about the
year 1850, but the ears, in correlation with its immaturity, are
relatively larger. The front view of the head of a male elephant
shot during (I believe) the sixties by H.R.H. the late Duke of
Saxe-Cobure- -Gotha in South Africa (text-fig. 107) seems un-
doubtedly to belong to the present type, the ears being of the
same large semielliptical form, and not rising, when cocked, above
the line of the middle of the head.
Text-fig. 108.
Head of Male Mashonaland Elephant (Elephas africanus selousi) in the
Imperial Institute.
Assuming that I am justified in separating the West Cape
Elephant, which survives only in a few protected localities such
as Mossel Bay, from 1. africanus capensis as typified by the Addo
Bush Elephant, the former will require a new name (unless
indeed it be H. africanus typicus). I accordingly suggest the
=
388 MR. R. LYDEKKER ON THE | Apr. 23,
name 1. africanus toxotis, taking the specimen shown in text-
fig. 106 as the type.
I now pass to the head of a male Elephant in the Inperial
Institute, shot by Mr. James Sligo Jameson. This Elephant (text-
fig. 108), as I learn from a footnote on page 433 of Mr. Selous’
‘A Hunter’s Wanderings in South Africa,’ was shot by Mr. Jame-
son in Matabililand (South Rhodesia).
Text-fig. 109.
Bupte pylons
tight Ear of Male Mashonaland Elephant (#/ephas africanus selousi),
from a specimen belonging to Mr. Selous.
Mr. Selous has kindly lent me the right ear of another Eleph:nt
(text-fig. 109) shot in Mashonaland (Rhodesia), which apparently
belonged to the same race as Mr. Jameson’s Elephant. ‘This ear
has now a vertical diameter of 4 feet 8 inches, but when fresh it
is stated to have measured 5 feet 6 inches. None of the upper
margin is now reflected, but in life about four inches appears to
have been turned back, as in H. a. knochenhaueri. The height of
the elephant to which it belonged was about 10 feet. The ear is
1907. | EARS OF AFRICAN ELEPHANTS. 389
much less elliptical than in #. a. foxotis, and approaches more to
that of the undermentioned /. a. cyclotis, but agrees with the
former in that the lappet underhangs part of the lower jaw and
chin.
J think I am justified in regarding the Mashonaland Elephant
as a distinct race, for which the name £. africanus selowsi would
be appropriate; the specimen in the Imperial Institute bemg
regarded as the type.
"This race will be characterised by the ears being of considerable
size, with the margin rounded, and the inferior lappet large,
pointed, and extending inwards to some extent beneath the lower
jaw and throat. In this respect they are unlike those of other
Kast African Elephants. Perhaps it may not be impertinent to
suggest to the authorities of the Imperial Institute that this
interesting specimen requires careful repairing at the hands of a
skilled taxidermist.
Text-fig. 110.
Head of Male Elephant from Swaziland, in the Imperial Institute.
A second head in the Imperial Institute (text-fig. 110) is stated
to have been obtained from Swaziland. The ears are in bad
condition, and I am unable to come to any certain conclusion as
to the race represented by this specimen, which appears, however,
to approximate to the Mashonaland type.
390 MR. R. LYDEKKER ON THE [Apr. 23,
The next specimen for consideration is the ear of the Congo
Elephant killed by Major Powell-Cotton (text-fig.111). The whole
contour is regularly rounded, and the transverse diameter rela-
tively large. That this type is quite different from the ear of
HK. africanus toxotis, as typified by the specimen represented in
text-fig. 106, 1s perfectly evident. Exclusive of the lappet, this
ear might well be described as oval; the lappet itself being broad,
blunt, and short. It thus accords in general character with the
Text-fig. 111.
Right Ear of the Congo Elephant (Hlephas africanus cyclotis °), from a male
killed by Major Powell-Cotton.
HL. africanus cyclotis of Dr. Matschie, from the South Cameruns ;
but in the absence of a figure of the type of the latter it is
impossible to say how close the resemblance really is. Under
these circumstances, all that can be done is to refer Major Cotton’s
specimen provisionally to the Cameruns race, with the suggestion
that if it prove distinct it should be named after the gallant
explorer.
The Congo Elephant, it may be added, differs from both the
190F. | EARS OF AFRICAN ELEPHANTS. 391
southern races in the large size of its tusks, which are, however,
frequently of a somewhat slender type.
Attention may now be directed to several photographs (text-figs.
112-116) of Elephant-heads from various parts of the eastern side of
Africa north of the Orange River, the ears of all of which are
characterised by the tendency to the assumption of a more o1 less
distinctly triangular form and by the pointed extremity of the
lappet, which is not inflected, and stands out quite distinct from
the side of the head. Dr. Matschie’s #. «a. knochenhaueri and
HE. a. oxyotis both pertain to this group; but whereas I have no
diticulty in identifying the latter type, it is less easy to decide
whether any one of my specimens should be identified with the
former, which is typically from German East Africa.
Text-fig. 112.
Head of a Male Elephant from North-west Rhodesia, in the pessession of the
Duke of Westminster.
1 take first the head in the possession of the Duke of West-
minster (text-fig. 112), which Iam told was killed by his Grace
in North-western Rhodesia. In this specimen the ears are of
medium size, rising, when cocked, slightly above the vertex of
the head, with the superior border nearly straight for some
392 MR. R, LYDEKKER ON THE [ Apr. 23,
distance, and the margin then forming a regular curve till the
width begins to narrow, when it runs obliquely to the extremity
of the small lappet which forms a symmetrical triangle. Although
LT have but little doubt that this specimen represents a distinct race,
I do not intend to propose a name, on account of its not being in
a well-known collection.
J now come to the male Hlephant trom Fort Manning, N.E.
Rhodesia (Nyasaland), mounted in the British Museum (Natural
History), of which the right side of the head is shown in text-fig.
113. The most striking fentumesof the ear in this specimen are its
relatively small size and distinctly triangular shape; the upper
border, which rises. considerably above the vertex of the skull
and is reflected posteriorly to a depth of several inches, being
Text-fig 113.
j
Buy re aworge
Head of Male Elephant from Fort Manning, N.E. Rhodesia (Hlephas africanus
knochenhaueri 2), from the entire specimen in the British Museum.
strongly arched and terminating in a marked angle which forms
the outermost point of the ear, while the lower and inner borders
form nearly straight lines meeting in the angular apex of the
lappet. The small relative size of the ear is indicated by the fact
that whereas the height of the specimen is 11 feet 4 inches, the
vertical diameter of the ear (inclusive of the reflection) is 4 feet
23 inches and the transverse diameter 3 feet 5 inches; the dimen-
sions of the ear thus being considerably inferior to those in the
Cape Elephant of 8 feet 8 inches in height recorded by Living-
stone. In addition to its small and triangular ears, the North-
east Rhodesian Klephant is also characterised by its relatively
small head and tusks. Although the dimensions and a figure of
1907. | EARS OF AFRICAN ELEPHANTS. 393
the ears of H. africanus knochenhaueri ave not available, the
Rhodesian animal clearly comes very close to that race (of which
the type specimen came from German East Africa), and is accord-
ingly, at any rate for the present, considered to be inseparable
therefrom.
Text-fig. 114.
Head of Male Elephant from the Aberdare Mountains, British East Africa (Hlephas
africanus peeli), from a complete specimen in the collection of Mr. C. V. A.
Peel at Oxford.
Next on the list comes an Elephant in the private collection
of Mr. C. V. A. Peel, 12 Woodstock Road, Oxford, shot in the
Aberdare Mountains, British Hast Africa, which are situated
immediately east of the Victoria Nyanza, at no great distance
from Mount Kenya. The ears of this specimen (text-fig. 114),
which are mounted “ cocked,” are remarkable for their length
and narrowness and somewhat pyriform shape; the vertical
diameter being 4 feet 33 inches and the transverse 2 feet 5 inches.
394 - MR. R. LYDEKKER ON THE [ Apr. 23,
The upper border is strongly arched, with the curvature continued
along the outer margin, where there is no angulation, for a
considerable distance, after which the ear rapidly narrows to the
extremity of the long and pointed lappet. The tusks are large
and of a relatively slender type. If I am right in identifying
the North-east Rhodesian Elephant with #. a. knochenhaueri,
it seems perfectly evident that Ma. Peel’s animal must represent
another race, which is equally distinct from the more northern
E. a. oxyotis.
Text-fig. 115.
Head of Male Klephant trom the Lake Rudolf Distmet (Hlephas africanus
cavendishi), shot by Ma. H. 8S. H. Cavendish, and now in the British Museum
(Natural History).
Since the above was written Mr. Rothschild has sent me a
photograph of the Klephant’s head from South-east Africa im
his museum, to which reference is made on page 282. It appears
in all respects similar to the head of Mr. Peel's spechnen, and
may be referred to the same race, of which the long slender
tusks form a feature. For this race I propose the name LZ. afr-
canus peeli, making Mr. Rothschild’s specimen a ‘ co-type.”
# t Krom the comparative nearness of the Lake Rudolf district to
the Aberdare Mountains, it might be reasonable to suppose that the
ROO EARS OF AFRICAN ELEPHANTS. 395
same type of Elephant would be found in both areas. The head
of an elephant from the former district, shot by Mr. H. 8. H.
Cavendish and now in the British Museum (text-fig. 115), appears,
however, to be distinct from Mr. Peel’s specimen, though the ears
are of the same general type in the two. Owing to the different
angles at which the photographs have been taken, it is difficult to
determine whether or no the ears of the present specimen rise so
high above the vertex as in Mr. Peel’s elephant. They are,
however, relatively broader (vertical diameter 2 feet 10 inches,
transverse diameter 2 feet 11 inches), and show a decided, although
not sharp, external angle, while the lappet is shorter and sharper.
They are considerably reflected at the sides, but scarcely at all at
the top. I think I may venture to make this specimen the type
of a race, with the designation EKlephas africanus cavendishi. It
will be distinguished from EB. a. knochenhaueri (as represented by
the British Museum specimen) by the larger and less distinctly
triangular ear, of which the upper border is more regularly
convex, the outer angle less defined, and the lappet Jonger and
narrower.
To #. a. cavendishi may be assigned the head of a bull Elephant
shot in the Galla Country, on the south-west border of Somali-
land, in the museum of Sir EK. G. Loder, to whom I am indebted
for the photograph of it. The Galla Country i is no great distance
from Lake Rudolf, and Sir E. G. Loder’s specimen is so like
the one given by Mr. Cavendish to the British Museum that
the photograph of the former might almost be mistaken for thet
of the latter. I think Mr. Gillett’s elephant also belongs to this
race.
Here I may refer to a very interesting photograph of a male
South Somali Elephant taken in the act of chare eing, with its ears
cocked, by Mr. R. McD. Hawker in 1906, and ‘presented by him
to the British Museum. The small scale of the photograph does
not admit of any definite statement with regard to this elephant,
which seems to have proportionately smaller ears than the fore-
going specimens.
Next on the list of races named on the evidence of the contour
of the ear comes the Sudan, or Abyssinian Elephant (A/ephas
africanus oxyotis) of Dr. M atschie, typically from the Upper
Atbara Valley, of which numerous examples have been recorded.
Dr. Matschie describes the ears as semicircular, with a very long
and sharp lappet. A very long and narrow triangle, of which
the upper side is convex, ter minating inferiorly in a long, sharp,
forwardly-directed la ppet, with the upper border reflected forwards
and the two ears overlapping in the middle line when in repose,
seems a better definition. It must, however, be admitted that
precise definition of the ear-characters of the various races is
almost an impossibility, and that reference to figures or specimens
is essential. With regard to the value to be attached to the
forward bending of the upper rim of the ear, Tam uncertain. In
many mounted specimens this rim is backwardly reflexed; but
396 MR. R. LYDEKKER ON THE [Apr. 23,
whether this 1s correct, or whether the condition alters according
as to whether the ears are in repose or cocked, | am unable to
state.
Text-fig. 116.
Head of the “ Queen’s Elephant,” an immature Male Sudan Elephant
(Elephas africanus ovyotis), formerly owned by the Society.
I take as a typical example of this race the head of the well-
known “ Queen’s Klephant” formerly in the Society’s collection
(text-fig. 116). This animal came from Abyssinia. A second
example of the same race is the young Abyssinian Elephant now
living in the menagerie in the Regent’s Park, whose ears accord
very closely in shape with those in the photograph of the “‘ Queen’s
Elephant.”
The largest ears I have seen are those of an Elephant killed
by Mr. A. Haig on the Blue Nile, one of which is shown in
text-fig. 117. This specimen, as mounted (possibly with some
stretching), has a maximum vertical diameter of no less than
1907.] EARS OF AFRICAN ELEPHANTS. 397
6 feet 54 inches, and a transverse diameter of 4 feet 14 inches.
The height of the animal to which it belonged is estimated by the
owner (from the circumference of the fore- foot) to have been over
13 feet. Mr. Haig also possesses the ears of another Elephant
from the same district, which are rather smaller. As mounted,
neither of these ears shows any fold at the top; although, as
already mentioned, in the “ Queen’s Elephant,” as well as in
‘Vext-fig. 117.
Right Har of a Male Sudan Elephant (Hlephas africanus oxyotis), from the
Blue Nile, in the possession of Mr. A. Haig.
the young Abyssinian Elephant now living in the Society’s
Gardens, there is a small forward fold. The enormous length
of the ears of this race is indicated in the photograph of a herd
at the Giza Zoological Gardens sent me by Captain §. 8. Flower,
where this feature is well displayed in one full-grown animal.
Despite the gigantic stature attained by #. a. oxyotis, the tusks
Proc. Zoou. Soc.—1907, No. XX VII, 270
398 MR, R. LYDEKKER ON THE [ Apr. 23,
in this race seem to be small and sharply curved; those of
Mr. Haig’s larger specimen weighing less than 60 lbs. each.
While the Elephant of the Blue Nile appears inseparable
from the Abyssinian form, i.e. 4. a. oxyotis, I am informed by
Mr. Haig that the Elephants of the White Nile are quite
distinct, having relatively small ears.
Texst-fig. 118,
Right Ear of the North Somali Elephant (H/ephas africanus orleansi), from the
type specimen in the collection of S.A.R. le Duce @’Orléans.
Whether the White Nile Elephant is or is not identical with
the one inhabiting Somaliland, I am unable to say; but the
head from the latter district belonging to the Duc dOrléans
(cf. text-fig. 118) clearly indicates a race markedly distinct. This
race, which I propose to call #. africanus orleansi (with the
type specimen in the collection of the Due d’Orléans at Wood
Norton), is characterised by the very small size of the ears, which |
do not reach within a considerable distance of the lower jaw and
throat, and are proportionately not much larger than those of
E. a. knochenhaueri, although of quite different shape. In the
1907. | EARS OF AFRICAN ELEPHANTS. 397
6 feet 53 inches, and a transverse diameter of 4 feet 14 inches.
The height of the animal to which it belonged is estimated by the
owner (from the circumference of the fore-foot) to have been over
13 feet. Mr. Haig also possesses the ears of another Elephant
from the same district, which are rather smaller. As mounted,
neither of these ears shows any fold at the top; although, as
already mentioned, in the ‘ Queen’s Elephant,” as well as in
Text-fig. 117.
Right Ear of a Male Sudan Elephant (Hlephas africanus oxyotis), from the
Blue Nile, in the possession of Mr. A. Haig.
the young Aby ssinian EHlephant now living in the Society’s
Gardens, there is a small forward fold. The enormous length
of the ears of this race is indicated in the photograph of a herd
at the Giza Zoological Gardens sent me by Captain 8. 8. Flower,
where this feature is well displayed in one full-grown animal.
Despite the gigantic stature attained by Z. a. ox yotis, the tusks
Proc. Zoot. Soc.—1907, No. XX VII, AT
398 MR. R. LYDEKKER ON THE [ Apr. 23,
in this race seem to be small and sharply curved; those of
Mr. Haig’s larger specimen weighing less than 60 lbs. each.
While the Elephant of the Blue Nile appears inseparable
from the Abyssinian form, i.e. H. a. oxyotis, I am informed by
Mr. Haig that the Elephants of the White Nile are quite
cistinet, having relatively small ears.
Text-fig. 118.
Right Ear of the North Somali Elephant (Elephas africanus orleansi), from the
type specimen in the collection of S.A.R. le Duc d’Orléans.
Whether the White Nile Elephant is or is not identical with
the one inhabiting Somaliland, I am unable to say; but the
head from the latter district belonging to the Duc d’Orléans
(cf. text-fig. 118) clearly indicates a race markedly distinct. This
race, which I propose to call H. africanus orleansi (with the
type specimen in the collection of the Duc d’Orléans at Wood
Norton), is characterised by the very small size of the ears, which
do not reach within a considerable distance of the lower jaw and
throat, and are proportionately not much larger than those of
E. a. knochenhaueri, although of quite different shape. In the
1907. ] EARS OF AFRICAN ELEPHANTS. 399
specimen figured, which indicates a small, although apparently
adult elephant, the maximum vertical diameter of the ear is only
2 feet 11 inches, and the transverse diameter 2 feet. The ears,
which (as mounted) show no folding, are remarkable for the
circumstance that their upper margin is almost continuous with
that of the head itself; the distinct notch occurring at the junction
of the ear with the head in all other Elephants that I have seen,
being absent. The upper margin of the ear forms a nearly straight
line inclining upwards to the outer upper angle; from the latter
point the outer margin runs nearly vertically dewnwards for a
considerable distance, and is then continued for a much longer
distance in a downward and inward direction to terminate in
the point of the inferior lappet. The lappet itself is small, tri-
angular, sharply pointed inferiorly, and separated by a wide notch
from the side of the lower jaw and throat. It seems, indeed, to
form a semi-distinct appendage of the ear, and is thus quite
unlike the corresponding element in any other head that has
come under my notice. The distinctness of the North Somali
Elephant, as it should be called, is thus perfectly apparent.
The next race for consideration is the one from the French
Sudan, typified by ‘“‘ Jumbo,” the Society’s well-known African
Elephant. ‘‘ Jumbo,” together with ‘‘ Sahib,” who died last year
after a sojourn of about thirty years at Paris, was brought to
France from some part of the French Sudan*, probably to the
southward of Lake Chad, and afterwards sold to our Society. At
the time he left England for America he was believed to be about
11 feet in height ; and both he and “Sahib” indicate an unusually
large race of the species. As regards the characters of the ear,
the West Sudan race appears to connect the Kast African races
with H#. a. cyclotis. Compared with that of H. a. oxyotis (text-
fig. 117) the ear of “Jumbo” (text-fig. 119) is relatively smaller,
and has a strongly emarginate, in place of a nearly straight
postero-inferior border. The emargination of this border causes
the lappet to be much more distinct from the rest of the ear than
is the case in H. a. oxyotis; and if this semidistinct lappet were
altogether removed, we should have an ear not very unlike that of
E. a. cyclotis, Jumbo’s ears, when in repose, nearly meet in the
middle line of the back, and show no flexure of the margin,
whereas the upper border of those of ZH. a. oxyotis is bent over
the front surface. A marked peculiarity in the case of “Jumbo”
is the deep channel running upwards and backwards from the
meatus. The subpyriform shape of the ear, with the above-
mentioned groove, and the absence of any flexure of the margin,
appear to be the most easily recognised features of the large West
Sudan race, which I propose to call /. a. rothschildi, taking the
statuette of “Jumbo” in the British Museum (Natural History)
as the type.
Thus ends this long review of the various forms of ear
* This information was given me by Mr. Rothschild after the paper was read.
21*
400 MR. R. LYDEKKER ON THE (Apr. 23,
presented by the Elephants that have come under my notice.
Possibly it may be pronounced an unsatisfactory one; and it is
confessedly but a preliminary recognisance, written in the hope
that it may lead to more definite results. Before such results can
be attained it is of prime importance that the British Museum
should obtain mounted heads of a number of Elephants from
different parts of Africa; and the assistance of sportsmen towards
this object is earnestly invoked, as if the work is to be done at
all, it is all important that it should not be delayed. IJ may also
express the hope that Dr. Matschie will speedily see his way to
publish photographs of the ears of the type specimens of the four
races recognised by himself.
Text-fig. 119.
Head of “Jumbo,” the male West Sudan Elephant formerly belonging to
the Society.
In regard to the dwarf Congo Hlephant (Hlephas africanus
pumilio) of Prof. Noack, this appears to be characterised, ac-
cording to our present information, primarily by its small stature,
which is estimated to be about 7 feet. There is, however, no
definite information even on this point, as the living specimen,
now in America, upon which the description was based is
immature. According to a note kindly communicated to me by
Dr. Chalmers Mitchell, this Elephant differs from the other races
of the species by the unusual shortness of the “finger” on the
1907. | EARS OF AFRICAN ELEPHANTS. 401
lower border of the tip of the trunk and unusual length of the
upper “finger.” It also appears to have a darker skin than in
EL. a. cyclotis. (See wmfra, p. 447.)
In conclusion, reference may be made to my account in the
‘Field’ newspaper for 1906 * of the skull of an Elephant killed
by Mr. Stanley C. Tomkins at the south end of the Albert Nyanza.
Mr. Tomkins, who presented the skull to the British Museum,
Text-fig. 120.
Front view of the Skull of the Sudan Elephant (Hlephas africanus oxyotis).
also forwarded a photograph of the dead elephant as it lay on its
side in scrub-jungle. Unfortunately the photograph does not
show the complete contour of the ear; but the portion visible,
although apparently of the triangular type, does not seem to
agree with the ear of any of the specimens referred to above.
According to information supplied by Mr. Tomkins, both the
natives of the Albert Nyanza and sportsmen who have visited
* Vol. vii. p. 1089 (1906).
402 MR. R. LYDEKKER ON THE | Apr. 23,
the district, are convinced of the distinctness and peculiarity of
the race of Elephants inhabiting that portion of forest country
lying to the south-west of the Albert Nyanza, in Congo territory.
Compared with a skull of #. africanus oxyotis presented to the
British Museum by the late Sir Samuel Baker (text-fig. 120), the
Albert Nyanza skull (text-fig. 121), while much inferior in total
length, is considerably wider across the forehead between the
temporal fosse. The forehead is also much flatter, the pit
between the sheaths for the tusks is shorter and shallower, and
the sheaths themselves are shorter and less divergent. There are
likewise marked differences in the occipital and lateral aspects of
the two skulls.
Text-fig. 121.
Front view of the Skull of the Albert Nyanza Elephant (Hlephas africanus
albertensis).
The skull is further remarkable for the great relative stoutness of
the zygomatic arch, which is absolutely thicker than in the larger
figured skull of #. a. oryotis, the large size of the infra-orbital
foramen, and the great thickness of the terminal alveolar portion
of the premaxille.
On one side of the upper jaw the two molars (penultimate and
last) are preserved, the former having the normal eight plates.
1907. | EARS OF AFRICAN ELEPHANTS, 403
Apart from the fact that all the other skulls of African
Elephants (some old and some young) in the British Museum are
of the same general type, such differences cannot be attributed
either to age or to individual variation. Indeed, if the Albert
Nyanza skull had been found in a fossil condition without the
teeth, there would be little hesitation on the part of naturalists
in regarding it as specifically distinct from the African Elephant.
The Albert Nyanza skull, it may be added, is quite different
from that of the Rhodesian Elephant referred to L. a knochen-
haueri, and apparently also differs as markedly from Dr. Matschie’s
description of that of the type specimen of the latter. The ear,
moreover, in Mr. Tomkins’s photograph is certainly different from
that of the Rhodesian Elephant in the British Museum. <A
small telephotograph of an elephant killed by Major Powell
Cotton on the Semliki side of the Albert Nyanza, recently pub-
lished in the ‘ Illustrated London News,’ may belong to this race,
and appears to have an altogether peculiar type of ear. Under
these circumstances the name of Hlephas africanus albertensis,
which I suggested in the ‘ Field ’ for the Albert Nyanza Elephant,
may stand.
The most noteworthy feature about the skull of /. a. albertensis
is its striking resemblance in contour to that of H. planifrons
of the Pliocene of the Siwalik Hills of Northern India, as will be
apparent by a comparison of the figure here given (text-fig. 121)
with the one of the fossil skull represented in plates ix. and x. of
Falconer and Cautley’s ‘ Fauna Antiqua Sivalensis.’ /. planifrons
is a member of the ‘ loxodont” group, with a ridge-formula very
similar to (although slightly higher than) that of H. africanus,
and one displaying primitive features in the development of suc-
cessional premolars.
Although accepting the view of the African origin of the Pro-
boscidea, I am of opinion (from the occurrence in those countries
of the intermediate “ stegodont” group) that the evolution of the
Mastodons into Elephants probably took place somewhere in
South-eastern Asia; and the similarity presented by the skull of
E. africanus albertensis to that of H. planifrons seems to suggest
that the African Elephant may be the descendant of the fossil
Indian species. The many remarkable instances of aftinity
between the Pliocene mammal!s of India and the modern mammals
of Africa (among which the relationship of Canis curvipalatus to
Otocyon is one of the most noteworthy), indicate that there is no
inherent improbability im such an ancestry for Hlephas africanus,
of which, on this view, /. a. albertensis is the most generalised
representative.
[ Postscript.—A second Elephant’s head in Mr. Rothschild’s.
museum, from some part of East Africa, appears to indicate an
animal nearly allied to the one from Swaziland of which the head
is represented in text-fig. 110 (p. 389). |
>
404 MR. OLDFIBLD THOMAS ON MAMMALS [ Apr. 23,
2. The Duke of Bedford’s Zoological Exploration in Hastern
Asia.—IV. List of small Mammals from the Islands of
Saghalien and Hokkaido. By OLprreLp Toomas, F.R.S.,
F.Z.8. (With Appendix on the Cold-blooded Verte-
brates, by G. A. BoutencEr, F.R.S., F.Z.8.)
[Received March 21, 1907.]
Whatever may be the riches of the St. Petersburg Museum in
collections from the far-eastern island of Saghalien, our own
National Museum has hitherto possessed scarcely a single mammal
from it, so that the results of a visit made to the island by
Mr. M. P. Anderson during last summer, presented to the Museum
by our President, are of the greatest value, and include examples
of a considerable number of species and subspecies new to science.
Mr. Anderson also paid a second visit to the island of Hokkaido
(formerly known as Yesso), and added a number of species to
those he had previously obtained there, a list of which was given
in my paper on his Japanese Mammals *.
The result of the present important collection is to show that
the two islands are exceedingly closely allied in their small
mammal faune. For nearly all the same species are represented in
both, although in the case of the Sciwrus, Micromys speciosus, and
Evotomys a slight difterence, here considered of subspecific value,
is perceptible. ‘The Saghalien species (apart from Bats) as yet
unrecorded from Hokkaido—Sorex daphenodon and minutus,
Sciuropterus russicus, and Sicista caudata—will probably yet turn
up in the more southern island. But the one Hokkaido species
absent from Saghalien, Micromys geisha, is so common and so
easily caught when present, that we may take it for granted that
it really does not occur in the latter island. Moreover, it should be
noted that the form of J. speciosws which occurs in Saghalien is
allied to the Korean peninsule, while that of Hokkaido is nearer
to the typical form found in Japan; so that in this genus, and
this genus only, Hokkaido would seem to be more allied to Hondo
than to Saghalien, a result probably due to the comparatively
recent immigration of the species concerned.
‘The forms found in each island are as follows :—
Saghalien.
Myotis mystacinus.
Sorex unguiculatus, daphenodon, shinto sevus,
minutus, gracillimus.
Sciuropterus russicus athene.
Sciurus vulgaris rupestris.
Tamias asiaticus.
Mus norvegicus.
* P, Z. S. 1905, i. p. 333.
1907.| FROM SAGHALIEN AND HOKKAIDO. 405
Micromys speciosus guliacus.
Craseomys bedfordie.
Evotomys anurensis.
Sicista caudata.
Lepus tumidus.
Hokkaido.
Rhinolophus ferrum-equinum nippon, cornutus.
Plecotus auritus.
Sorex unguiculatus, shinto sevus.
Sciurus vulgaris orientis.
Tamias “ lineatus.”
Mus norvegicus.
Micromys speciosus ainu, geisha hokkaido.
Craseomys bedfordie.
Evotomys mikado.
Lepus tumidus ainw.
The present collection consists of 341 specimens, belonging to
22, species and subspecies, about half coming from each of the
two islands referred to.
Of previous publications on the subject, reference need only be
made to the following work :—
V erte-
Nixousky, A. M.—Survey of Saghalien and its Fauna.
brate Animals. St. Petersburg, 1889.
35 mammals are recorded, but they are mostly large species, of
which Mr. Anderson was unable to obtain any examples.
The following are some extracts from Mr. Anderson’s notes.
It may be observed that he was the first person, not a Japanese,
to enter the island of Saghalien after the cession of its southern
portion to Japan.
“The island of Saghalien has a length of nearly 600 miles by at
most 120 miles in breadth. In the north it is separated from the
mainland by a shallow strait only 5 miles in breadth, while its
southernmost point is 25 miles from the northern part of
Hokkaido.
“Tts two southern peninsulasare the continuations of mountain
ranges leading up into the centre of the island, where they
approach each other, but do not joi. Korsakoff, the capital, is
situated at the head of the gulf between the peninsulas.
‘Both mountain ranges of southern Saghalien and the unculti-
vated parts of the valley are densely forested with coniferous trees
(chiefly larch and fir), among which are mixed large numbers ot
birches, alders, and elms. The flora, indeed, is strikingly like
that of Hokkaido. Heavy dews and frequent rains keep the soil
constantly moist.
“ My first collecting place (July 11th to 25th) lay 15 miles N.W.
of Korsakoff, on the bank of a river which, draining a portion of
the main range, empties into Aniwa Bay at its N.W.edge. Here
406 MR. OLDFIELD THOMAS ON MAMMALS [ Apr. 23,
we were in the level valley, forested at this spot chiefly with larch,
but, in such clearings as existed, growing rank with many herbs.
“ From July 25th to August 5th I spent ata point some 7 miles
up the same river. Here we were at the foot of the mountains of
the main range, but we found few mammals on the forested hills.
The best collecting was in the clearings, and open spots in the
forest where grass, reeds, and ‘ weeds’ grow rank.
“ August 10th I began work at Dariné, and remained there till
August 28th. Dariné is 25 miles N.W. of Korsakoff, and stands in
the valley at the edge of the western mountains. Here, as else-
where, we found both forest and clearings.
“‘T spent two days in the mountains west of Dariné, at an
altitude of 1000 feet. Here the musk-deer is said to be plentiful,
but I saw no evidence of it besides some scraps of skin and a long
tooth shown me by a Japanese settler. The single chipmunk
(Tamias) and other specimens were secured at this place, but
these are all found in the lowlands as well.
Notes on places visited in Hokkaido.
“From Sept. 7th to Sept. 21st I worked at Ochiai, a village in
nearly the exact centre of Hokkaido. This place is at an altitude
of 1300 feet, and situated on a tributary of the Ishikari River, a
little west of the crest of mountains that run from south to north
through Hokkaido. The region is of steep though not lofty
mountains, well covered with their native forest, which is largely
coniferous in character. During my stay the weather proved
pleasant, but chilly. This is said to be the coldest part of the
island in winter.
“‘T spent the time from Sept. 24th to Oct. 6th at Kuchan, best
described as 30 miles S.W. of Sapporo. This place is at the foot
of a mountain called Shiribeshiyama, an extinct volcano closely
resembling Fujiyama, though not so high. My collecting was
done in the forest of oaks, chestnuts, maples, and elms that covers
the foot of this mountain and the neighbouring hills. Altitude
900 feet or below.”
1. RHINOLOPHUS FERRUM-EQUINUM NIPPON Temm.
Ss. 1113. Kuchan, 30 miles 8.W. of Sapporo, Hokkaido,
2. RHINOLOPHUS CoRNUTUS Temm.
6. 1118. Kuchan, Hokkaido.
The occurrence of these two species of Rhinolophus appears to
be the most northern record of any Leaf-nosed Bat in the Hast, but
in the warmer West, of course, they range considerably further
north.
3. Puecorus Aauritus L.
3g. 1028. 2. 1027. Ochiai, Central Hokkaido.
Although the Long-eared Bat has often been said to occur in
1907.] FROM SAGHALIEN AND HOKKAIDO. 407
Japan, these are the first examples from the Far Hast that the
British Museum has received. They show remarkably little
difference from Kuropean examples.
4. Myoris Mysracinus Leisl.
3. 875, 880. 2. 874, 876. 17 miles N.W. of Korsakoff,
Saghalien.
3. 923. $. 951. Dariné, 25 miles N. of Korsakoff.
5. SOREX UNGUICULATUS Dobs.
3. 816, 877, 878. ©. 817. 17 miles N.W. of Korsakoff,
Saghalien. 150’.
6. 946. Mountains 35 miles N.W. of Korsakoff. 1000’.
Ce Doe rood Olan Io0n) Darme, 2onmnlesmN Nem Or
Korsakoff. 200’.
6. 961, 1026. 2. 960, 990, 1031, 1032, 1047, 1056. Ochiai,
Central Hokkaido. 1200’.
g. 1097, 1111. Kuchan, 30 miles 8.W. of Sapporo, Hokkaido.
This large-footed Shrew was discovered in Saghalien by Dr. L.
von Schrenk, by whom the type, afterwards described by Dobson,
was sent to the St. Petersburg Museum. No example of it had
hitherto reached the British Museum.
No. 899, a female, is smaller than usual, with a smaller skull,
shorter tail, and lighter claws ; but I can find no sufficient reason
for distinguishing it from other specimens taken at the same
place. Mr. Anderson notes of this individual that it ‘ contained
5 embryos, 9°5 mm. long ; mammee 6 in number, all inguinal.”
6. SOREX DAPHHNODON, sp. 0.
©. 916, 925, 956. Dariné, 25 miles N.W. of Korsakoff,
Saghalien.
S. araneus group. The teeth very heavily pigmented; tail
thickly haired.
Size and general characters of S. araneus. Hairs of back
about 43 mm. in length (summer specimens). General colour
above dark brown (between Prout’s brown and histre). Sides
little hghter than back; under surface dull greyish, with slight
drabby tinge. Handsrather heavy, though not nearly so large as
in S. unguiculatus, the claws slightly longer than usual; upper
surface of hands and feet grey-brown, darker than in S. annexus.
Tail of medium length, heavily haired and pencilled even in.
summer specimens; the hairs at the tip over 7 mm. in length.
Skull and dentition very much as in S. araneus, except that the
teeth are extraordinarily heavily pigmented throughout, the
pigment covering all the cones of the teeth and passing lower
into the valleys than in any Sorex known to me; the hypocones
of the upper p*, m', and m’° all heavily pigmented ; viewed exter-
nally the limiting line of the brown pigment is more than halfway
down the outer side of the teeth.
408 MR. OLDFIELD THOMAS ON MAMMALS [ Apr. 23,
Dimensions of the type, measured in the flesh :—
Head and body 59 mm. ; tail 38; hind foot (s. u.) 12°5; ear 8.
Skull—condylo-basal length 18°5 mm.; basal length 15-7;
greatest breadth 9°6; greatest breadth across molars 5; vertical
height from basion 5-9; front of i’ to back of m? 8-2.
Type. Adult female. B.M. No. 7.2.5.16. Original number 916.
Collected 17th August, 1906.
This Shrew is distinguishable from all other Eastern species
known to me by its heavily pigmented teeth. In general appear-
ance it is very like S. annexus Thos., but besides the difference in
the teeth, its more hairy tail and differently coloured teeth at
once distinguish it.
I owe to the kindness of Dr. Allen examples representing his
S. buaxtoni, described from Gichika Ochotsk Sea, and find that it
has a smaller and slenderer skull (greatest breadth in type 8 mm.)
and that its teeth are not more heavily pigmented than usual.
7. SOREX SHINTO SHVUS, subsp. n.
6. 787, 850. 15 & 17 miles N.W. of Korsakoff, Saghalien.
6. 948. ©. 947. Dariné, 25 miles N.W. of Korsakoff.
Ss. 1030, 1064. ©. 995, 1050. Ochiai, Central Hokkaido.
6. 1112. @. 1096. Kuchan, 30 miles S.W. of Sapporo,
Hokkaido.
Quite like the true S. shinto Thos., of Hondo, in all respects,
but larger throughout, except that the tail is only of about the
same length.
The following are the external dimensions of four specimens,
measured in the flesh :—
Head &body. Tail. Hind foot. Ear.
mm. mm. mm. mim.
Saghahien. <¢. 787 (Type). 69 50 12°5 8
us ORO AE eee 58 48 1295) 8
Igloo, 5 IW gcebee 62 49 13 8
4 Oh CIOS euaanheye ti 58 50 12°5 8
Skull (of type)—greatest length 18°6 mm.; basal length 16:3 ;
greatest breadth 9-2; length of upper tooth-series 7:8.
Hab. Both Saghalien and Hokkaido. Type from 15 miles
N.W. of Korsakoff, Saghalien.
Type. Old male. B.M. No. 7.2.5.19. Original number 787.
Collected 15th July, 1906.
This Shrew is evidently of the same type as the Hondo S. shinto,
discovered previously by My. Anderson, but differs, as most
northern forms so often do, by having a larger body, with a
proportionally shorter tail. I can find no difference whatever
between the Saghalien and Hokkaido specimens.
8. SOREX MINUTUS GRACILLIMUS, subsp. n.
@. 921. Dariné, 25 miles N.W. of Korsakoff, Saghalien. 200’.
18th August, 1906. B.M. No. 7.2.5.23. Type.
1907. | FROM SAGHALIEN AND HOKKAIDO, 409
Size and proportions as in the smaller forms of European
minutus. Skull very light and delicate, peculiarly narrowed in
the facial region. In ordinary minutus the skull narrows evenly
forward from the brain-case, but in gracillimus, while the brain-
case is of about the usual breadth, the narrowing is much more
abrupt in the interorbital region and the whole face in front of
this is particularly light and slender. P* and molars rather
narrower than in true minutus. Pigmentation of teeth slight,
but the specimen is old, and they are all somewhat worn.
Dimensions of the type, measured in the flesh :—
Head and body 51 mm.; tail 44; hind foot 11; ear 6.
Skull—condylo-basal length 15°2 mm.; basal length 13:8;
greatest breadth 7-7; interorbital breadth at hinder end of ante-
orbital foramina 2°8; length of upper tooth-row 6:8; breadth
across molars 3-4.
Hab. & Type as above.
Of this tiny Shrew, the Eastern representative of our Pygmy
Shrew, Mr. Anderson only sent the skull and one hind foot, the
rest having been stolen by a rat. Fortunately the measurements
were preserved, and as these and the skull are the chief basis for
systematic work on Shrews, the colours being comparatively
unimportant, I have thought myself justified in giving a name to
the animal on the very perceptible difference in the shape of the
skull. No Far-Eastern forms of this group have been previously
recorded, except under the old names of Si) minutus or pygmeus.
SCIUROPTERUS RUSSICUS * ATHENE, subsp. n.
2. 865, 17 miles N.W. of Korsakoff, Saghalien, Ist August,
1906. B.M. No. 7.2.5.24. Type.
Colour as in S. momongat ; skull as in true russicus.
General colour above drab-grey, the tips of the hairs clay-
colour; under surface dull whitish, the sides slightly washed
with reddish brown. Upper surface of hands and feet smoky
grey, becoming black on the digits ; the hairs at the bases of the
claws clay-colour ; hairy part of soles greyish white; lower side
of digits naked. Tail very much as in S. momonga, the upper
and under layers of hair black, the middle layer isabella or clay
colour, the hairs at the tip verging towards buffy.
Skull with the long palatal ‘foramina and large bulle of
S. russicus, each of these parts being markedly smaller in
S. momonga.
* Pteromys russicus Tiedem. Zool. p 451, 1808. American zoologists have
rightly shown that the name volans Linn. belongs to the American species; but
this is noé because Linnzeus’s Mus volans is founded on Seba’s Pl. xliv. fig. 3 3,
“ exclusively American,’ as the animal figured by Seba is clearly a Petaur ista, in
spite of his statement that its locality was Virginia. The real basis of the Linnean
name is Ray’s Sciurus americanus volans (Quadr. p. 215, 1693), which is undoubtedly
the American species.
+ Or at least as in S. momonga amygdali Thos., the restricted momonga being
only represented by a bleached specimen.
410 MR. OLDFIELD THOMAS ON MAMMALS [ Apr. 23,
Dimensions of the type (immature) :—
Head and body 125 mm.; tail 110; hind foot 36:5; ear 20°5.
Skull—ereatest length 37 mm.; basilar length 28; zygomatic
breadth 22; interorbital breadth 7; palatilar length 16-2;
palatal foramina 4°8; length of bulle 10:1; length of upper
tooth-series 72.
Hab. & Type as above,
This Flying-Squirrel seems to be a local race of the Russian
and Siberian S. russicus, rather than any relation of S. momonga,
though its colour is exactly as in the latter, instead of the clearer
grey of its ally. Satunin’s S. bwechnert*, from Kansu, has a
shorter hind foot and a more rufous colour.
10. ScruRUS VULGARIS RUPESTRIS 7, subsp. n.
@. 812. 15 miles N.W. of Korsakoff, Saghalien. Sea-level.
3. 927. ©. 952-958. Dariné, 25 miles N.W. of Korsakoff.
200’.
g. 944. 9. 945. Mts. 35 miles N.W. of Korsakoff. 300’ &
1000’.
Quite like S. v. orientis, in summer pelage, but markedly
smaller throughout. Winter pelage not represented.
Colour in summer as in orientis, but there appears to be a
greater proportion of melanism, for not one of the six specimens
is unaffected in this way. The least affected (No. 927, the type)
has the feet and flanks blackish, while the only one (No. 812)
which has the feet and flanks reddish, has the back of a very
darker colour. The back of the type is rather redder than
‘“‘ Prout’s brown.”
Skull decidedly smaller than in orientis, 3-4 mm. less in
extreme length, and the three true molars about half a millimetre
_less in combined length. This last point is important, as several
of the specimens are immature, so that their full size can only be
gauged by their teeth.
Dimensions of the type (slightly immature) :—
Head and body 190 mm.; tail 174; hind foot 55; ear 32.
Skull—greatest length 48°8 mm.; basilar length 37; length of
true molar series 6°7.
Dimensions of No. 944, a fully adult male :-—
Head and body 208 mm. ; tail 178; hind foot 57; ear 32.
Skull—egreatest length 50:7 mm.; basilar length 39°5; length
of nasals 14:3; breadth of brain-case 24; palatilar length 22°5 ;
combined length of p* and three true molars 8°9.
Type. Immature male. B.M. No. 7.2.5.26. Original number
927. Collected 20th August, 1906.
In a group where the main distinctions have been made on
colour, it is somewhat embarrassing to have to deal with this
* Ann. Mus. St. Pétersb. vii. 1902, p. 3 (1903).
+ Iam informed by Dr. Knud Andersen that Saghalien is a modification of a
Chinese word meaning “cliff at (the mouth of) Amur.”
1907.] FROM SAGHALIEN AND HOKKAIDO. 411
Squirrel, which differs in size alone from its Hokkaido ally, but
there seems no doubt that it ought to have a name. Possibly
some colour character will be found when winter specimens are
available for comparison.
11. ScrurRUS VULGARIS ORIENTIS Thos.
3. 1065, 1073. Ochiai, Centra) Hokkaido.
g. 1077, 1103. @. 1075,1110. Kuchan, Hokkaido.
12. Tamras asraticus Gmel.
Q. 949 (immature). Mts. 35 miles N.W. of Korsakoff,
Saghalien.
13. Tamtas “ LINEATUS Siebold.”
3S. 1039, 1063. Ochiai, Central Hokkaido.
2. 989, 1006, 1019, 1038, 1049, 1070, 1071, 1072. Kuchan,
Hokkaido.
This fine series, representing the first mammal ever described
from Japan, “ Myoxus lineatus” Siebold*, is of much value to
us, but for want of more Siberian material [am unable tc venture
an opinion as to its specific or subspecific relationship to 7.
asiaticus. But if not identical, it is certainly very closely allied
to the latter.
14. Mus norvecicus Erxl.
3. 811. 15 miles N.W. of Korsakoff, Saghalien.
15. Micromys SPECIOSUS GILIACUS, subsp. n.
54 specimens (mostly immature) from 15 miles N.W. of Korsa-
koff (sea-level), 17 miles N.W. of Korsakoff (150'), Dariné (200’),
and mountains 35 miles N.W. of Korsakoff (1000’).
Most closely allied to the Korean subspecies, J/. s. pen-
insule +, with which it agrees in the proportionally long and hairy
tail, as compared with the Japanese forms, but distinguished by
the ear being uniformly shorter. In the considerable number
of peninsulee examined, the ear is measured by Mr. Anderson as
15, 15:5, or 16 mm., generally the last in adult specimens. In the
Saghalien form, on the other hand, the great majority have the
ear measured as 14 mm., some two or three only being labelled
as 15.
The colour of the Saghalien specimens is rather darker than in
those from Korea, but the former are all in summer and the
latter in winter pelage, which may account for the difference.
The fur, in summer pelage, is not so distinctly spinous as it is
in Japanese specimens, but there is a certain crispness which may
develop into spininess in old age.
* Spic. Faun. Japon., in Diss. Hist. Nat. Japon. p. 13, 1824.
+ P. ZS. 1906, p. 862.
412 MR. OLDFIELD THOMAS ON MAMMALS [ Apr. 23,
Detailed measurements :—
Head and body. ‘Tail. Hind foot. Kar.
mM. mim. min, mm.
6. 881 (Type). 104 110 24 14-5
Cie SSUES on -oioe 104 hl 24 14
OF B82 ise ae 103 113 23°D 14
Obs BOM ose ese 102 108 22 15
Skull of type—greatest length 28 mm.; basilar length 21:8 ;
zygomatic breadth 13-7 ; interorbital breadth 4°6; palate length
13°1; diastema 9; palatal foramina 5°7; length of upper molar
series 3°7.
Type. Adult male. B.M. No. 7.2.5.42, Original number 881.
Collected at Dariné, 11th ios 1906.
This is no doubt the “ Jus sylvaticus” of Nikolsky’s work, and
indeed it is probable that all the Long-tailed Field Mice that
have been recorded from the Far East of Asia belong to this or
some closely allied form, with 2—2=8 mamme, and that none
of them are really members of the J/. sylvaticus group, with
1—2=6 mamme.
The following is a short synopsis of the Japanese and Korean
forms of Micromys, which have been referred to or described in
the present series of papers :—
A. Size large. Hind foot (s.u.) over 21 mm. and skull
over 25. Supraorbital edges ridged.
a. en not lineated .. aunpoupacgaseescenses Ll SDCCIOSIIS:
Tail shorter than head and body.
ey Foot large, 26-28 mm. in adults. Hokkaido ... M. s. ainu.
t3. Foot medium, 22-24 mm. in adults.
a‘. Tail generally over 100 mm. Hondo &c....... M. s. speciosus.
64. Tail 90 mm. or less. Okils. ..................... Ms. navigator.
62. Tail ionger than head and body.
c3. Ears longer, 15-16 mm. Korea ..................... MW. s. peninsule.
d3. Kars shorter, 14-15 mm. Saghalien ............... M. s. giliacus.
6. Back with a more or less distinct median line.
IKGGREAN ais. eaten steerer SARS SR een Sauna PE UEME IVER CUO MOLIPZOUSS
B. Size smaller. Hind foot under 21 mm. and skull
under 25. Supraorbital edges rounded .................. WL. geisha.
c. Ears smaller, averaging about 18 mm. Saghalien...... M. g. hokkaidi.
d. Bars larger, generally about 14 mm.
c?. Size larger, tail longer. Head and body 85-95 mm.
Tail 90-100 mm.
e8, Hind foot shorter, 18-20 mm. Hondo &c.......... M. g. geisha.
ae 3, Hind foot longer, 20-21 mm. Yakushima.. . Meg. yakui.
. Size smaller, tail shorter. Head and body about
gOmm. Tail 80mm. Oki Is. ....0......... M. g. celatus.
16. Micromys sPEctosus ArInuU Thos.
18 from Ochiai, Central Hokkaido, and Kuchan, 30 miles
S.W. of Sapporo.
17. Micromys GEISHA HOKKAIDI Thos.
51 from Ochiai and Kuchan, Hokkaido.
1907. ] FROM SAGHALIEN AND HOKKAIDO. 413
18. CRASEOMYS BEDFORDIZ Thos.
33 from all the localities visited in Saghalien.
19 from Ochiai, Central Hokkaido. 1200’.
jo L090 2 LOMGs Kuchany S0gmiless sain or Sapporo:
Hokkaido.
I can find no appreciable difference between the Saghahen and
Hokkaido specimens of Craseomys.
19. KvoroMys AMURENSIS Schrenk.
59 specimens from Saghalien (all localities visited).
The type specimen of this species was obtained by Schrenk
close to the mouth of the Amur, and I have no doubt that the
present series are referable to the same form. It is probably
the “ Arvicola rutilus” of Nikolsky.
From /. mikado, of Hokkaido, Schrenk’s species is distinguish-
able by having the pale lateral area rising up much higher over
the shoulders, so as to narrow the reddish upper colour to an
ill-defined band little more than half an inch broad along the
nape and withers. In #. mikado the shoulder area is rarely
lighter or higher than the general light lateral colour. In
addition the staal, or at least the brain- case, of mikado seems to
average rather longer and narrower than in amurensis, though
the difference is not very great, and the two forms are no doubt
very closely allied.
20. Evoromys mrkabo Thos.
29 from Ochiai, Central Hokkaido. 1200’.
21. SICISTA CAUDATA, Sp. n.
©. 862. 17 miles N.W. of Korsakoff, Saghalien. 150’.
Ist August, 1906. B.M. No. 7.2.5.104. Type.
‘“¢ Among tall grass."—U/. P. A.
A species without dorsal stripe, and with a very long brown
unicolor tail.
Size rather larger than in S. concolor Biichn. Fur close and
fine; hairs of back, in a summer specimen, about 6 mm. in
length. General colour along the dorsal area pale brown, of a
rather warmer tone than Ridgway’s “ wood-brown,” the sides
paler and with a slight buffy suffusion. Under surface pale
greyish with a drab suffusion, line of demarcation on sides not
strongly marked. Visible part of ears (proectote and metentote)
well-haired, dark brown. Upper surface of hands silvery white,
of feet similar except that the metatarsals are dusky mesially
for their prominent third. Tail very long, well-haired, uniformly
brown above and below. Mamme 2—2=8.
Skull lightly built, smoothly rounded. A deep concavity
present in the single specimen on the median frontal suture just
behind the nasals; perhaps not normal. Palate very much as in
S. leathemi, the edge of the mesopterygoid fossa close behind the
raised ridge which runs across the palate between the last molars,
Proc. Zoou. Soc.—1907, No. XX VITI. 28
414 ON MAMMALS FROM SAGHALIEN AND HOKKAIDO. [ Apr. 23,
and distinctly in front of the anterior limit of the parapterygoid
fossee ; the latter much narrower anteriorly than in S. tianshanica.
Dimensions of the type, measured in the flesh :—
Head and body 63 mm.; tail 115; hind foot 18; ear 14:5.
Skull—ereatest length 20°77 mm.; basilar length 15°3; zygo-
matic breadth 10°6; interorbital breadth 4; breadth of brain-
case 10; palatilar length 8:5; palatal foramina 4:1; length of
upper cheek-tooth series (crowns) 2°7.
Hab. & Type as above.
The discovery of the genus S%cista in Saghalien is a considerable
extension of its known range, as the furthest eastern point from
which it appears to have been recorded is Kan-su, W. China.
Sicista caudata seems to be most nearly allied to S. concolor
Biichn. from Kan-su, as S. subtilis is at once distinguishable by
its dorsal stripe and elongated palate, and S. Jeathemi Thos. and
tianshanica Salensky by their bicolor tails. From S. concolor it
may be separated by its longer tail and shorter tooth-series,
while other differences will no doubt be found when examples
of the two forms can be directly compared.
22. Lepus tTimipus L.
3 (young). 924. Dariné, 25 miles N.W. of Korsakoff,
ee
On the Cold-blooded Vertebrata of Saghalien.
By G. A. Boutencer, F.R.S.
One Lizard, Lacerta vivipara, one Snake, Vipera berus, and
one Frog, Rana temporaria, species existing over the whole of
Northern Europe and Asia, were the only known representatives of
the Reptiles and Batrachians hitherto recorded from Saghalien *.
The collection made on the island, 15-25 miles N.W. of
Korsakoff, by Mr. Malcolm P. Anderson in J uly and August
1906, and presented to the British Museum by His Grace the
Duke of Bedford, includes several examples of the Common
Lizard, and representatives of the following species :—
Tropidonotus vibakari Boie—Manchuria, Japan.
Anceistrodon blomhoffi Boie—KE. Siberia, China, Japan.
Rana amurensis Blgr.—Manchuria, N. China.
Rana esculenta L., var. chinensis Osb.—Corea to Siam,
Japan.
Bufo vulgaris Laur.—Kurope and N.W. Africa to Manchuria,
China, and Japan.
Bombinator orientalis Blgr.—Manchuria, N. China.
The single fish collected by My. Anderson is referable to
Gastrosteus steindachneri Jordan & Snyder.
* Of. A. M. Nikolsky’s work (in Russian) on the Vertebrates of Saghalien,
(St. Petersburg, 1889).
1907. ] ON NEW WORMS OF THE FAMILY EUDRILIDA, 415
3. On some new Species of Harthworms of the Family
Eudrilide, belonging to the Genera Polytoreutus, Neu-
mannella, and Eminoscolex, from Mt. Ruwenzori. By
FRANK EH. Bepparp, M.A., F.R.S.
[Received April 9, 1907.]
(Text-figures 122-127.)
The following pages relate to a number of species of Eudrilide
which I received from the Natural History Museum through the
kindness of Mr. W. R. Ogilvie-Grant, F.Z.8. They were collected
upon Mt. Ruwenzori along with a number of other species of
Oligocheta belonging to the genera Benhamia and Alma. These
latter genera have been lately investigated by Signor Cognetti de
Martiis*, upon material collected by H.R.H. the Duke of the
Abruzzi trom the same locality. I have therefore limited myself
to the description of the Hudrilidee, of which specimens must, I
should presume, have been collected by the Italian expedition ;
but they have not, so far as I am aware, up to the present been
described. All the species are new, but are referable to genera
already defined, which genera are in every case Hast African in
range.
Polytoreutus ruwenzorii, sp. n.
Of this species the collection contained but a single example,
and that in a not very good state of preservation for dissection,
I have, however, been able to ascertain, as I think without doubt,
that the species is new and allied to a small group of species of
this genus of which all the members known hitherto have been
described by Michaelsen’?. This group—which includes the species
P. kirimaensis, P. usindjaensis, and P. sylvestris—is limited to the
shores of Victoria Nyanza, Albert Nyanza, and the neighbouring
country; and the occurrence therefore of an ally upon Mt.
Ruwenzori is not surprising. The likeness of these four forms is
to be seen chiefly in the peculiar relations of the diverticula of the
spermathecal pouch and, in three of them at any rate t, in the
existence of paired copulatory pouches debouching to the exterior
on either side, and independent, of the penis. The present species,
whose exact locality within this area I fix by means of its specific
name, 1s represented by an example quite fully mature which
measures 77 mm. in length by 5-6 mm. in breadth. It is there-
fore a stoutish but also shortish worm.
The sete ot Polytoreutus ruwenzorti are disposed like those of
other species of the genus: 7. e. the ventral setee are much wider
apart than the lateral sete. The distance between each seta of
the ventral pair is something like three times that which separates
Boll. Mus. Zool. Torino, vol. xxi. notes i. and iii. ; vol. xxii. note xiv.
“ Regenwiirmer,” in Deutsch-Ost-Afrika, 1896.
Apparently not in P. usindjaensis.
toh &
IQ*
416 MR. F. E. BEDDARD ON NEW [Apr. 23,
the individual sete of the lateral pair. I have endeavoured
to make an exact study of the distribution of the sete upon
the clitellar segments, concerning which there is some but
not exhaustive information, of some other species of the genus
already described; for this character seems to be one of probably
systematic value. On these segments I could only find one of the
two sete of the lateral pair, and the seta present was the inner-
most. I ought to mention that these statements depend upon a
microscopic examination of the entire cuticle stripped from the
body, and not merely upon an inspection of the entire worm with
alens. The apertures through which the sete are protruded are
so obvious that the failure to find one is strong evidence of its
absence. The ventral setz, on the other hand, were present apon
the clitellar segments with the exception of the xviith, where only
the outer seta of the pair was present. Ventrally the clitellum is
not so strongly developed as it is laterally and dorsally, which facts
may be related to the presence or absence of sete.
The clitellum ot Polytoreuius ruwenzorii is, like that of Poly-
toreutus sylvestris and some, but not all, other species, best
developed laterally and dorsally. Ventrally it is not so well-
developed, and here the intersegmental furrows are plainer than
laterally. It embraces segments xiii. to xvil., which is the usual
extent of the clitellum in this genus.
The nephridiopores lie in front of the lateral pair of sete, in
front of each pair, and not in front of any one seta of the pair
more particularly. They commence apparently in the fourth
segment. A notable fact with reference to these pores is that
when the cuticle is stripped off—and I have mapped the pores by
this means—-a considerable strip of the (as it would therefore
appear) chitinous lining of the duct of the nephridium is also
stripped off and protrudes from each aperture. I have not
noticed anything of this kind in other Oligocheta.
The oviducal pores are quite conspicuous and lie upon the xivth
segment behind and to the outside of the nephridial row and
the lateral seta of that segment. The single male pore is on the
border of segments Xvi. [xviii and the sper mathecal | pore behind it
upon the interval xviii. / RIBS
The internal anatomy of this species, so far as concerns the
alimentary and circulatory organs, seems to agree with that of the
next species to be described and with the members of this genus
generally.
The sperm-sacs are like those of Polytoreutus generally (but not
P. bettonianus) in being exceedingly long, and at their commence-
ment and for a long way back of much less diameter than they
ave more posteriorly. The sacs extend for more than 30 segments
back from their point of origin. That of the right side is fifteen
segments longer than the shorter sperm-sac of the left side. The
difference in length in this species is more pronounced than in
that next to be described. The dilated chambers at the beginning
of the sperm-duct immediately after it leaves the don ell are
1907. ] WORMS OF THE FAMILY EBUDRILIDS. A417
conspicuous in this as in the next species. The spermiducal
glands are peculiar in form and do not altogether agree with those
of P. sylvestris, to which they appear to come nearer in structure
than to those of other species of the genus Polytoreutus. They agree,
however, with the last-named species in the fact that the duct of
the gland instead of emerging, as is the rule among these worms,
from the end of the spermiducal gland, leaves the gland some
little way in front of the proximal end. Hach gland is rather
bent in form, but otherwise lies straight. It is of firm consistency,
but is not covered with a sheath of muscle appreciable to the naked
eye or through a lens. The slight bending of the corresponding
glands in Polytoreutus sylvestris figured by Michaelsen is rather
exaggerated in the present species. They appear also to be rather
longer in P. sylvestris than in P. ruwenzorii. Michaelsen does
not mention in that species a character which is very noteworthy
in P. rwwenzorii. He describes the “ prostate” glands indeed
merely as being “‘unregelmiissig eingeschniirte.” In the specimen
of P. rwwenzorti reported upon here the surface of the gland was
much marked by furrows, and the appearance given was that of a
very long gland tightly coiled up with ‘some concrescence between
the individual loops of the coil. There is no indication of any-
thing of the kind in the figures given by Michaelsen either of
P. sylvestris or of its allies. The two copulatory chambers
mentioned by Michaelsen* in P. sylvestris and P. kirimaensis are
quite as large in P. rwwenzorw as in those species.
As in several species—for example, Polytoreutus kirimaensis T,—
the present species of Polytoreutus is to be characterised by a
very slender spermathecal sac which lies beneath the nerve-cord,
than which it is no thicker. It is thus difficult to see, and, as
Michaelsen has remarked, is apt to escape the eye. Particularly
was this the case with the worm described in the present
communication. For the contents were very slight in certain
regions of the sac, which rendered it even more ditlicult of obser-
vation. It is certainly no wider than the nerve-cord, which
overlies it. It is largely by virtue of the different forms which
the spermathecal sac shows in this genus that the species of
Polytoreutus are discriminated.
The species which I name Polytoreutus ruwenzorw is quite
different in details, so far as concerns this organ, from any other
species of which descriptions have been published. It comes
nearest to Polytoreutust kirimaensis so far as I can gather, but
shows obvious differences from that species.
The median spermathecal sac is slender as in that species
and is straight or nearly so in its course beneath the nerve-cord,
not much convoluted as in the allied Polytoreuius sylvestris §.
* Toc. cit.
+ Michaelsen, “ Die Regenwtirmer Ost-Afrikas,’ in Deutsch-Ost-Afrika, vol. iv.
1896, p. 16. { Loe. cit. pl. i. fig. 21.
§ Where, however, it is also occasionally less convoluted, perhaps in less mature
individuals (Michaelsen, Joc. cit. pl. ii. fig. 23).
418 MR. F. E. BEDDARD ON NEW [Apr. 23,
Anteriorly the sac passes into the fourteenth segment near to the
anterior wall of that segment without any change. Arrived here it
ends in two diverticula of short extent. These diverticula are appa-
rently of much shorter extent than in any of the species Polytoreutus
kirimaensis, P. usindjaensis, and P. sylvestris, whose spermathecal
apparatus is built upon the same plan as that of P. ruwenzorii.
Michaelsen, as a matter of fact, does not differentiate, except in
the case of P. usindjaensis, between each diverticulum and the
oviduct with which it becomes continuous, which in fact opens into
it according to my interpretation of these various structures in the
genus Polytoreutus*. In the figure annexed hereto this arrange-
ment is rendered plain. The sudden diminution of the cecum of
Text-fig. 122
2
Spermathecal sac of Polytoreutus ruwenzorii.
d. Diverticulum of sac. o.d. Oviducal pore. 2. Spermathecal pore.
the spermathecal sac (text-fig. 122) where it is continuous with the
oviduct is obvious. Moreover, the oviduct is extremely long as
compared with that of some other species, and is much coiled.
Much more so is this the case with Polytoreutus ruwenzoru than
with either of the three species mentioned as coming nearest to it
in respect of the spermathecal sac and its forward diverticula.
But apparently these three species do agree with P. rwwenzorti in
having a much longer oviduct than in many other species of the
genus. There is a further point of agreement between the new
species described in the present paper and the three Hast-African
species with which I have compared it in the nature of the
* P. Z. S. 1902, vol. 1. p. 206 et seq.
1907. | WORMS OF THE FAMILY EUDRILIDA. 419
spermathecal diverticula. The slender spermatheca just in front
of the terminal section of the male efferent apparatus divides into
two branches, which diverge at right angles and run dorsally, this
portion of the spermatheca forming a tube more than 5 mm. im
length. Towards the dorsal median line the tube of each side
turns back upon itself after emitting a very short but shghtly
swollen diverticulum ; the recurrent branch runs alongside of
the outgoing branch, ‘the two forming a loop which suggests at
first sight a nephridium. It is indeed not very much thicker
than the nephridia. The returning branch then, having arrived
at the level of the point whence it departed, dorsally turns at
right angles and joins its fellow a little in front of the external
aperture, which is quite inconspicuous. This peculiar origin of
the diverticulum on each side is exactly matched in the “three
species to which the present is evidently allied, viz. Polytoreutus
usindjaensis, P, kirimaensis, and P. sylvestris. But in all of the
three species mentioned the diverticulum itself is of some
considerable calibre. Polytoreutus kirimaensis comes nearest tc
the present species in that the diverticulum is smaller than in the
other two. But even in this species it is much larger than in
P. ruwenzorii. There is thus no ditticulty in discriminating
Polytoreutus ruwenzorii by the structure of the female efferent
and copulatory apparatus. It is very interesting to notice that
Polytoreutus ruwenzorti also shows points of resemblance to P.
sylvestris and P. kirimaensis (but apparently not to P. wsindjaensis)
in the structure of the male efferent apparatus, which has been
already described. There seems to be no doubt that these
four species form a little group of Polytoreutws; but it is not
possible in my opinion to separate them off from the other species
as a genus or even a subgenus, at least at present.
It is clear from the above account that Polytoreutus ruwenzorii
comes nearest to P. sylvestris of Michaelsen. It is only, as I think,
with this species that we have to reckon in determining the dis-
tinctness or otherwise of the Polytoreutus which I regard as new.
The most plain differences from this species are to be seen in
the small size of the spermathecal diverticula and the very
reduced length in front of the undivided portion of the posteriorly
fused spermathecal sacs. It may be thus defined :—
Polytoreutus ruwenzorii.
Length 70-80 mm.; breadth 4-5 mm. Distance between sete
of ventral pair three times that between sete of lateral pair.
Outer seta of lateral pair absent on clitellar segments. Clitellum
saddle-shaped. Male pore xvit./xviii.; spermathecal pore aviii./xin.
No genital area behind pores. Spermathecal sac bifurcate for a
short distance in front; posterior diverticula very small ; oviduct
between spermathecal sac and receptaculum very long. Right
sperm-sac longer than left. Spermiducal glands giving off duct in
front of proximal end gland, muth furrowed. Copulatory chambers
present,
420 MR. F. E. BEDDARD ON NEW [Apr. 23,
Polytoreutus granti, sp. n.
In describing some years since* several species of this genus
from Hast Africa, I found among a collection from Mt. Kenya
two closely allied species, which, however, were plainly to be
differentiated upon a careful study. Itis interesting to find upon
Ruwenzori the samepresence of two closely allied species of
Polytoreutus, not—it may be remarked—specially related to their
congeners of Kenya. To find closely related species in the
same comparatively restricted area is rather more remarkable
than would have been the existence of more vemotely allied
examples of the same genus. This species, which I have named
after Mr. Ogilvie-Grant, F.Z.S., comes nearer to Polytoreutus
kirimaensis than does P. ruwenzorii. It is represented by a
single specimen, not fully mature as to the clitellum, but appa-
rently quite fully mature as to the sexual organs. One of the
two copulatory chambers and the penis were protruded. The
size and the external characters generally agree with those of
P. rwwenzorii.
The worm is a trifle more slender. The clitellum was not
developed, and upon the segments to be included in it I observed
no deficiency of setee such as occurs in P. rwwenzorii. The re-
lations between the distances which separate the two sete of each
pair are much asin P. ruwenzorii. In the same way I observed a
long tube of chitin to be extruded from the nephridiopores. I do
not like to assert positively that there is a difference between
the two species in the segment which contains the first pair of
nephridiopores. But in the present species I noted a pair of
these apertures in the third segment, 7. e. a segment further
forwards than I observed the same pores in P. ruwenzorii. The
internal anatomy seems to agree with that of P. ruwenzorii and
other species of Polytoreutus in the alimentary canal with its
appended calciferous glands and in the situation of the last
heart (eleventh segment). It may be mentioned, however, that
P. granti, like P. rwwenzorti, has the dorsal vessel doubled
in the twelfth segment. This doubling of the dorsal vessel ig
known in the genus Polytoreutws—tor example, in P. gregorianus *.
The male organs of reproduction are much like those of
P. ruwenzorii, and yet show differences in minutie. As in that
and other species of the genus, there is but a single vas deferens
on each side, ending in front in an elongated chamber (‘‘Samen-
magazine”) behind the funnel. The sperm-sacs are but a single
pair. They are elongated and not so markedly thin anteriorly as
in P. ruwwenzorii and other species. The right-hand sac, as in
that species,is longer than the left, but the difference is not
quite so pronounced. The length of the longer sac is 21 mm.
* “ On some new Species of Earthworms belonging to the Genus Polytoreutus, &c.,”
P. ZS. 1902, vol. ii. p. 190.
+ Beddard, P. Z.S. 1901, vol. i. p. 191. Michaelsen has not referred to the con-
dition of the dorsal vessel in the species with which the present is particularly
compared.
1907. | WORMS OF THE FAMILY EUDRILID&. 42]
The two sacs are not joined at their distal extremity. The
terminal apparatus of the male efferent ducts is quite like that
of P. ruwenzorti. In precisely the same way (text-fig. 123) the
spermiducal gland is almond-shaped, and somewhat bent upon
itself at the point whence its duct emerges; the surface is not,
however, quite so strongly furrowed. The copulatory chambers
seem to be exactly as in P. rawwenzorit.
Terminal male organs of Polytoreutus granti.
c.c. Copulatory chamber. p. Spermiducal gland.
The female organs of generation (text-fig. 124), on the other hand,
show greater differences from the same organs in P. ruwenzori.
There is the same slender median spermathecal sac which underlies
the nerve-cord and is hardly convoluted in its course. Nor is it
of any greater diameter than the nerve-cord. Anteriorly this sac
divides into two, but there is no marked division near to the
point of bifurcation of the sac between the spermathecal sac
and the oviduct which opens into it. This break is very clear
in P. rwwenzorii. And in that species the diverticula* of the
spermathecal sac are short, the greater part of the coiled tube
intervening between the unpaired spermathecal sac and the
receptaculum being the oviduct. In the present species I could
not ascertain the precise spot where the oviduct debouched into
the diverticula of the spermathecal sac; but this point is at any
rate very far removed from the point of bifurcation of the sperma-
thecal sac; the greater part of the coiled tube, therefore, which
intervenes between the unpaired spermathecal sac and the re-
ceptaculum being referable to the diverticula of the spermathecal
sac. This important difference between these two species, other-
wise very nearly allied, isremarkable. It is apparently correlated
with another structural feature in which they differ. In examining
* The word “diverticulum ” is, of course, not strictly correct. The two spermathece
are fused in the middle and separate at both ends.
422 MR. F, E. BEDDARD ON NEW | Apr. 23,
microscopically this part of the reproductive apparatus in ely-
cerine after removal from the body, I noted in addition to the
receptaculum, called by Michaelsen the “ Hitrichterblase,” a sphe-
rical chamber which obviously corresponds to what Michaelsen
calls the ‘‘ Ovarialblase,” and which is believed by him to contain
the ovary. In Polytoreutus cerulews * this Ovarialblase forms the
end of the branch of the spermathecal sac into which it opens on
the one hand, and is connected on the other with a narrow tube
communicating with the Hitrichterblase, which is, I think, part
of the funnel of the oviduct pulled out by the retreating sperma-
thecal sac. J cannot be certain that the spherical chamber in
Polytoreutus granti has the same connections, but I cannot help
regarding it as the homologous structure. I have not represented
it in the figure (text-fig. 124), Now this appears to be wanting
in Pols ytoreutus ruwenzori altogether, as it is, for example, in
Polytoreutus magilensis. Where it does occur it appears to
mark the boundary between the oviduct and the spermathecal
sac; if so, then the present species has, as I have already sug-
gested, a very long diverticulum to the spermathecal sac and a
short oviduct *.
Text-fig. 124.
t
Spermathecal sac of Polytoreutus granti.
Lettering as in text-fig, 122.
Polytoreutus granti differs from its ally P. ruwenzorii in
possessing, like P. hirimaensis and P. sylvestris, considerable
appendages to the spermathecal sac posteriorly. These measured
in my example 6 mm., and were therefore Just a trifle shorter
than the spermiducal glands, which measure 7 mm. in length. The
proportions, in fact, are not very different from those of P. syl-
vestris ; but in the present species the length of both structures
* Michaelsen, JB. Hamb. wiss. Anst. ix, Taf. iv. fig. 30.
+ Beddard, Quart. Journ. Micr. Sci. n.s. vol. xxxiv. pl. xxv. fig.
ie
1907. | WORMS OF THE FAMILY EUDRILIDE. 423
seems to be considerably less than in P. sylvestris, where they
extend very much further back in the body. As in that species,
however, the spermathecal diverticulum on each side receives or
emits the duct leading to the exterior from the side and not from
the end. The relationships of the diverticulum to the thread-
like regions of the spermathecal sac which enter and leave it were
precisely like those depicted by Michaelsen for P. sylvestris, and
not like those of P. kirimaensis; for in the latter species the
spermathecal diverticulum simply bifureates at its end into the
incurrent and excurrent regions of the spermathecal sac.
The above account of the anatomy of Polytoreutus granti
shows that it cannot be confused either with the species which
I have just described or with any other known form. It
comes nearest to P. ruwenzori and to P. sylvestris. It differs
most markedly from P. rwwenzorii by the characters of the
spermathecal diverticula, and by the great length of the anterior
undivided portion of the spermathecal sac. It differs from
P. sylvestris mainly in the presence of a well-marked circular
chamber at the end of the undivided spermathecal sacs, and by
the shorter spermathecal diverticula and spermiducal gland. It
may be thus defined :—
Polytoreutus granti.
Length 70-80 mm.,; breadth 4-5 mm. Distance between sete
of ventral pair three times that between sete of lateral pair. Male
pore xvii./aviii., spermathecal pore aviit./aix. Spermathecal sacs
long and much coiled in undivided anterior region; posterior
diverticula of some length. Oviduct between spermathecal sacs and
receptaculum ovorum not long. Right sperm-sac longer than left.
Spermiducal glands furrowed ; duct arising before proximal end of
gland. Copulatory chambers present.
Neumanniella ruwenzorii, sp. n.
IT refer two fully mature, moderately large individuals, as well
as a number of smaller specimens, of an earthworm, apparently
new to zoology, to this genus Vewmanniella*, for reasons which
the following account of its structure will render plain. The
principal distinguishing feature of the genus is thus described by
its founder, viz.:—‘‘ Die fiir mehrere neue Arten aufgestellte
Gattung Mewmanniella unterscheidet sich von den verwandten
Gattungen Hminoscolex, Gardullaria und Telendrilus durch die
vollstindige Unpaarigkeit der Samentasche.” This is plainly to
be seen in Vewmanniella ruwwenzorii. The larger of the two
examples is 105 mm. long and measures 3 mm. in diameter. It
is not strongly pigmented. The prostomiwm is very small and
retracted within the peristomial segment. :
The sete have the usual arrangement met with in this genus.
The individual setz of the ventral pair are much wider apart
* Michaelsen, “ Die Oligocheten Nordost-Afrikas,’ Zool. Jabrb. (Abth. f. Syst.)
xvii. p. 501.
424 MR. F, E, BEDDARD ON NEW [ Apr. 23,
than the closely paired sete of the lateral pair. The distance
between each seta of the ventral pair is fully five times as great
as that which separates the two sete of the lateral pair. The
sete are rather small. On some segments, at any rate, of the
chtellum there are no setz present at all. This is certainly the
case with segment xiv., where the exact position of the lateral
seta between the nephridiopore in front and the ovipore behind
could be easily fixed. There is no trace upon the cuticle (which
was stripped off and examined) of these sete or of the orifices
through which they protrude. In NVewmanniella siphonocheeta
Michaelsen particularly notes that sete are present* upon the
clitellum. Nothing is said upon the matter in the case of other
species. ‘The absence of setze upon the clitellum is well known to
occur in certain species of Pheretima, while other species have them
on the clitellum.
The nephridiopores ave very plainly visible upon the clitellar
segments only with the use of a lens; but they are not thus visible
upon the other segments of the body. ‘The reason for this is not
wholly, if at all, the turgescence of the clitellar segments, which
thus makes the pores obvious. When the cuticle is stripped off
it is very distinctly to be noted that the pores themselves are
smaller in size upon the preclitellar than upon the clitellar
segments. The difference is very considerable. This can hardly
be the result of stretching, and must indicatea larger nephridium,
or, at least, a larger terminal duct to the nephridium. I observed
the first nephridiopore upon the third segment. Michaelsen states
(of the species t where he notes the point) that the nephridiopores
hie in line with the pair of sete cd. I found in Newmaniniella
ruwenzorw a decided relationship to seta c. These pores are, 1t
should be added, near to the anterior dividing-line of their
segment.
The clitellum of Newmanniella ruwenzorti is complete all round
the body. It is as strongly developed upon the ventral as upon
the dorsal side. Its yellow colour contrasts with the rest of the
body. The clitellum shows some variation from species to: species
of this genus in the fact of being saddle-shaped or as in the
present species. It begins upon the xuith segment, the posterior
one-third or so of which is invaded by the clitellar epithelium.
At the other extremity it ends upon the xvilith segment, so that
the dimensions are as in other species of the genus, and as in the
Kudrilide generally. I have already remarked upon the apparent
absence of sete upon this region of the body; there are also no
traces to be observed externally of penial sete in the neighbourhood
of the male generative pore or elsewhere.
The most anterior of the g generative pores is the spermathecal
aperture. his is very conspicuous in the middle line and just
on the boundary-line of segments xiii. and xiv. It is rather a
small orifice, but nevertheless quite evident. There is no modi-
* Loe. cit. p. 502.
+ NV. siphonocheta and N. tenuis, loc. cit. pp. 502 & 505.
1907. | WORMS OF THE FAMILY EUDRILID®. 425
fication of the integument in its neighbourhood. On the xivth
segment are the paired orifices of the oviducts. ‘These lie exactly
behind the nephridiopores of that segment and not far from the
posterior boundary of the xivth segment. Theapertures are quite
conspicuous, but not much (if at all) larger than the nephridio-
pores upon the clitellum, which, as already mentioned, are wider
than are those apertures upon the segments in front of the clitellum.
The male pore is a single aperture like the spermathecal pore.
It lies exactly on the boundary-line of segments xvii. and xviii.
It is considerably larger than the spermathecal pore, and the
actual orifice is surrounded by an area having a rather transparent
appearance. There is no protrusion of the Bursa propulsoria, nor
is the orifice situated upon an area which is at all raised beyond
the general level of the body. Apart from the slightly modified
integument surrounding the male pore, which has no counterpart
in the case of the spermathecal pore, the body of this species shows
no genital papille.
With regard to internal structures I have no observations to
make, save those based upon examination with a lens and a micro-
scope in the case of detached pieces of certain organs and systems.
I have not investigated this Eudrilid by means of sections on
account of its poor state of preservation.
With regard to the alimentary tract, the paired calciferous
glands seem to me to be rather further back than the thirteenth
segment, but as the worm was much softened I should not like
to be quite certain. The condition of the proximal swellings
upon the sperm-ducts seems in this genus to offer systematic
characters for the differentiation of the species. For in JV. tenis
these ‘‘Samenmagazine” are hardly marked at all, and they
are quite conspicuous in iV. siphonocheta. In the present
species these structures are present, but rather different in
their condition from those of some other Hudrilide. Each of
these swellings upon the sperm-duct is of oval form, tapering
towards each end; but instead of lying immediately after the
funnel of the sperm-duct, there is a considerable stretch of narrow
tubular sperm-duct before the funnel. The latter lies deep within
the sperm-sac. | have observed the characters of these bodies in
the smaller specimens, not in that whose structure has served
for the preparation of the rest of the present description of the
species.
Vhe terminal male efferent apparatus is constituted as follows :—
Each of the two prostate or spermiducal glands measured about
10 mm. in length, and each gland was folded only once upon
itself, thus forming a U. The diameter of each gland was not
more than, if indeed quite so much as, 1 mm. The (morpho-
logically) posterior end of each gland lay further forwards in the
body than the proximal or anterior end of each spermiducal
gland. Anteriorly each tubular spermiducal gland suddenly
narrowed to form a firm and slender duct; the two ducts running
backwards soon join and form an unpaired tube which constitutes
426 MR. F, E. BEDDARD ON NEW [Apr. 23,
one limb of a U, the anteriorly directed limb being the Bursa
propulsoria. The spermiducal glands have a a soft opaque appear-
ance; they are not hard and with a nacreous glitter as in some
Eudrilids. On the other hand, the Bursa propulsoria has an
almost metallic appearance to the naked eye. It is slender and
fusiform. There are no penial sete.
The spermathecal gland of this Eudriliid was unfortunately cut
into when the animal was slit open along the back. It is more
or less globular in shape, and occupies about the first half of the
clitellum. It is quite dorsal in position, lying immediately beneath
the body-wall, and has pushed the dorsal blood-vessel to the
left. The contents were an opaque white granular mass, in which
Text-fig. 125.
j 2
e
d 1
0.0.
Spermathecal apparatus of Newnanniella ruwenzorit.
d. Diverticula of spermathecal sac (8). Funnel of oviduct. o0.d. Ovidueal pore.
® , Spermathecal pore.
could be detected nothing resembling a spermatophore, though
the friable coagulated matter may have been the material out of
which aspermatophore was to be constructed. The spermathecal
sac (text-fig. 125), it should be said, widens out from its duct,
which lies to the right side of the gut. I could see no indication
of any communication between the spermathecal sac and the
cavity of the intestine, such as occurs, for example, in Parascolen.
The narrow duct of the spermathecal sac shows no nacreous
glitter, nor does it ultimately widen out into any structure that
may be termed a Bursa copulatrix. On the contrary, the duct of
IO WORMS OF THE FAMILY EUDRILIDS. 427
the spermathecal sac just before its opening on to the exterior
suddenly narrows to half its former calibre and becomes more
muscular, with transverse and longitudinal fibres. At the point
where this narrowing begins, two ceca, one on each side, are
given off and slightly curled backwards. These arise therefore
from what I have termed the duct of the spermatheca. These
cca are rather longer than the very narrow terminal chamber of
the spermatheca, with a diameter half again the width of that
chamber; each cecum is about half the diameter of the wider
part of the duct of the spermatheca. From the extremity of each
cecum arises a short tube with very weak muscular walls; this
becomes a little wider, and at a short distance from the sperma-
thecal ceecum contains the ovidueal funnel, which can be seen to
fan out within the chamber so formed. ‘The oviduct apparently
also opens partly into the receptaculum ovorum ; the short oviduct
opens on to the exterior in the usual way. The receptaculum
ovorum was full of eggs. I could find no ovary within the
chamber into which the funnel of the oviduct opens, and suspect
that the ovarial tissue has been entirely transferred to the
receptaculum ovorum. In any case, the various sacs and ducts
mentioned appeared to form a closed system in which the ovary,
if persistent elsewhere, was not included. I am not clear how
far this species agrees and disagrees with others of the genus.
In three of the four species described by Michaelsen, that naturalist
figures a tube which encloses the oviducal funnel at one end and
opens at the other into the spermathecal sac. There is no in-
dication of any diverticula of the spermathecal sac which receives
the ‘“‘ Verbindungsschlauch,” such as I find in NVeumanniella
ruwenzoru. Nor could I, as already mentioned, detect a special
ovarian sac lodging the ovary and communicating with the rest
of the egg- conducting apparatus, such as Michaelsen found. It
should be observed that this bifurcation of the spermathecal
sac anteriorly to receive the oviducts is exactly like the disposition
of this sac in Polytoreutus. Its presence in the species Vewman-
miella ruwenzorti necessitates a revision of the generic characters
used by Michaelsen, who uses as a generic character the fact that
“Samentasche ganz unpaarig.” This character alone therefore
serves to discriminate the present species from all of those
described by Michaelsen. It may be thus defined :—
Neumanniella ruwenzorii.
Length 105 mm., breadth 3 mm. Distance between ventral sete
about five times that between dorsal sete. Some of the sete absent
from clitellum. Clatellum complete, xiii./aviti. Male pore single,
median, upon xvi.faviu. Spermathecal pore single, median, upon
wit. / cw. Spermiducal glands with well-marked narrow long duct,
each of which joins its fellow 0 open into fusiform muscular
terminal chamber. No penial sete. Spermathecal sac with two
diverticula, which receive oviducts at extremities, and a small narrow
muscular Bursa propulsoric.
428 MR. F. E, BEDDARD ON NEW [ Apr. 23,
Eminoscolex ruwenzorii, sp. v.
I refer to a new species of the genus Hminoscolex a worm
which is considerably softened, but in which the more important
characters are nevertheless plainly visible. The completely paired
condition of the male and female organs, coupled with the ventral
calciferous pouches in segments ix., X., xi. and the paired glands
in segment xiii., are decisive of its generic position.
Ventral view of Hminoscolex ruwenzorit.
Some of the segments are numbered.
The worm measures rather more than 200 mm. in length by
4-5 mm. in width, and is thus the largest species of the genus.
The colour above is dark purplish brown. The clitellum was not
developed. The seé@ are paired; but the phrase descriptive of
these structures in Michaelsen’s amended definition of the genus*,
viz., “ Borsten ventral sehr weit, lateral enger gepaart,” hardly
applies to the present species.
* “Die Oligocheten Nordost-Afrikas,” Zool. Jahrb. (Abth. f. Syst.) Bd. xvii.
p. 482.
1907, ] WORMS OF THE FAMILY EUDRILIDA. 429
The ventral sete (text-fig. 126) are rather wide, much as in Poly-
toreutus, but the lateral sete are only slightly more approximated,
the proportions being about 5:6. It isremarkable that on the nine
or ten anterior segments of the body the ventral sete are very much
larger, perhaps twice the size of the lateral sete; this discrepancy
ceases after that point, and both pairs of sete are equisized and
small. The ventral pair of sete of the xviith segment are
absent.
The nephridiopores lie in front of the lateral pair of sete, and
are not specially related to one or other of the pair. They appear
to begin in the third segment.
The oviducal pores are upon the xivth segment, in line with the
nephridiopores.
The spermathecal pores ave paired as in other species of the
genus. The pore of each side lies in front of the outer of the
ventral pair of sete. The pores are not very large.
On the following intersegmental furrow (i. e., xili./xiv.) are
two pairs of minute orifices, which lie on a level with each of the
four ventral sete. They are rendered more conspicuous by being
surrounded with a yellowish area, This area is the expression of
internal sacs, which probably correspond to the copulatory glands
found in many EHarthworms of the families Megascolicide and
Geoscolicide, but not, I believe, hitherto recorded among the
Eudrilide. These sacs, though small, are very easily to be seen
when the septum separating segments xili. and xiv. is pushed
forward.
The male pores are very large and conspicuous, and lie between
segments Xvil. and xviil. in a straight line with the spermathecal
pores. The flaps of skin surrounding the pores cause them to be
rather obliquely set, as is shown in the accompanying figure
(text-fig. 126). The hinder margin of each orifice is much
thickened and forms a glandular pad; the two very nearly meet
in the middle line behind.
The intersegmental septa are not very much thickened, and
they cease to be at all thickened after that which separates
segments xi./xil. The gizzard, which is well developed, though
not very large, lies undoubtedly in segment vy. The median
calciferous pouches are in segments 1x., x., and xi. The paired
calciferous glands of segment xiii. have rather an unusual form.
Kach consists of a relatively narrow tubular coiled structure very
much longer than it is broad. This les coiled on each side of the
gut in the xiith segment.
The dorsal blood-vessel is nowhere double; the last pair of
hearts is In segment x1.
The male organs are much like those of other species of this
genus. The present species is holandrous, and the sperm-sacs are
two pairs, more or less tongue-shaped bodies, in segments xi.
and xii. The ends of the sperm-ducts near to where they open
into the sperm-sacs are, as in other Eudrilide, dilated into what
Michaelsen has termed a “Samenmagazin.” As in other species
Proc. Zoou. Soc.—1907, No. X XTX. 29
430 ON NEW WORMS OF THE FAMILY EUDRILIDZ. _[ Apr. 23,
of Hminoscolex, the region of the sperm-ducts in question is not
only widened but is of some length and coiled, forming a body of
oval contour constituted by the closely approximated windings of
the tube. It is conspicuous through its white colour as in other
Eudrilide. The sperm-duct opens into the tip of the spermiducal
gland (text-fig. 127), which is in this species directed forwards ; the
entire gland is sausage-shaped and bent once upon itself. It ends
in a narrow muscular duct which opens into the rather large
Bursa propulsoria; the latter is of circular contour. There are
no penial sete.
Text-fig. 127.
Terminal male organs of Hminoscolex ruwenzorit.
B.c. Bursa propulsoria. p. Spermiducal gland. v.d. Vas deferens.
The female organs of generation were so much softened that
some details have probably escaped me. It is, however, plain
that there are a pair of nearly spherical spermathecal sacs which
are perfectly free from each other and do not communicate, as in
some species of the genus, at the distal extremity with each other.
The oviduct is very long and straight in its course ; anteriorly it
is seen to communicate with a rather small receptaculum ovorum,
which lies close to the septum dividing the xilith from the xivth
segment to the outside of the oviduct. In the other side is a slight
swelling which appears to be fixed against the septum dividing
the same segments, which doubtless represents the ‘“ Hitrichter-
blase” and the “ Ovarialblase.” I am doubtful whether this
chamber is also in communication with the spermathecal sac
or surrounds it. It is not, however, necessary to settle this
point) for purposes of the identification of the species, which T
believe is sufficiently distinguished from other species of the genus
hitherto described. Of these there are ten species known.
A peculiarity of Hminoscole ruwenzorti is the thickened fold
1907. ] MR. H. SCHERREN ON HYBRID BEARS. 431
which bounds the male pores posteriorly, and is comparable to a
rudimentary penis or pair of penes. Of this structure there seem
to be the equivalents in a few other species, viz. in 1’. sylvestris ™,
EL. barnimit, H. newmannit, EH. viridescens, and perhaps
EH. toreutus§. Although the mere thickening which I have
figured in #. rwwenzorii is different from the projecting penis of
E. viridescens, the immaturity of my specimen might account for
the difference very easily.
But Eminoscolex ruwenzorii is to be distinguished from LZ. syl-
vestris and H. barnimi by the fact that the dorsal setz are further
apart than in those species, where the distance a—6 is three times
the distance c-d. It cannot be confused with #. newmanne by
reason of the fact that in the latter species the spermathecal
pores are opposite te the lateral sete, whereas in L. rawen-
zorit they are, as in the majority of species, in front of the
ventral sete. There remain /. toreutus and LZ. viridescens. In
the latter species ||, however, the spermathecal pores are a segment
further back and are placed in a common depression ; in Z. toreutus
the spermiducal glands appear to have a different form and the
spermathece are larger. Hminoscolex ruwenzorii may be thus
defined :—
Eminoscolex ruwenzorii.
Length 200 mm. ; breadth 4-5 mm. Ventral sete rather further
apart than lateral, the proportions being 6:5. Ventral setw of nine
anterior segments enlarged. Male pores avii./xviii., with thickened
posterior margins (a penis?) ; female pores xii./xiti., in line with
seta b. Copulatory-gland pores on xitt./xw. Proximal ends of
four sperm-ducts widened and twisted into a closely adpressed coil.
Spermiducal gland of only moderate length, with narrow muscular
duct. Spermathece more or less spherical, not conjoined.
4. Some Notes on Hybrid Bears.
By Henry ScHERREN, F.Z.S.
| Received April 9, 1907. |
The fact that some months ago the Hon. Walter Rothschild
purchased from Stuttgart and deposited in the Society’s Gardens
two hybrid Bears attracted attention to these animals, and
induced me to look into the literature on the subject. Bears play
a considerable part in folk-lore; and one might have imagined
there would be many references to cases of hybridity, seeing that
these animals have been credited with carrying off women. The
following story is quoted by Gesner §[ from Gillius :—
‘““De Philippo Cosseo Constantiensium sacris prefecto, ingenu-
* Michaeisen, Zool. Jahrb. (Abth. f. Syst.) xviii. p. 486.
+ Td. MT. Hamb. wiss. Anst. xvii. L Id. ibid. xiv.
§ Id. “ Regenwiirmer,” in Deutsch-Ost-Afrika, p. 9.
|| Michaelsen, Oligocheta in ‘ Das Thierreich,’ Lief. 10 (Berlin, 1900), p. 407.
€ ‘ Historie Animalium,’ lib. i., p. 1068.
Zoe
432 MR. H. SCHERREN ON HYBRID BEARS. [Apr. 23,
arum artium perstudioso, qui hoc ipsum se ex bono authore
cognitum habere, mihi valde affirmavit: ursum accepi ex montibus
Allobrogum puellam in speluncam rapuisse, eandemque venereo
complexu et osculatione prosecutum fuisse: atque ex pomis
agrestibus, que permulta quotidie in speluncam inferret maturiora
studiose delegisse, eidemque edenda amatorie dedisse, ac nimirum
cum ad cibi inquisitionem proficisceretur, iIngenti saxo speluncam,
ne puella exire posset, occludisse. Cum autem post longam
inquisitionem parentes ursinum latibulum preterirent, suam
peradolescentem animadvertisse, saxoque egre depulso, eam
recepisse.”
Similar stories have been told from remote antiquity of the
anthropoid apes, and among them cases of hybridity, though
suspected, are not established. I am acquainted with no case
of hybridity in Bears in a wild state, and with but few in
zoological collections. Dr. Brandes has suggested that the
absence of records of wild hybrids is due to the fact that the
species occupy regions widely separated; and that the paucity of
menagerie-bred hybrids may be accounted for by the fact that
there is always the fear that putting two individuals in one cage
may have bad results, since a fight between two adult bears is
sure to terminate fatally for at least one of the combatants.
In an article in the ‘English Cyclopedia’ Broderip refers to
a belief in hybrids between the dog and the bear; and says that
an account of such a creature was given in the ‘ Histoires
Prodigieuses extraictes de Plusieurs Fameux Auteurs Grecs et
Latins, sacrez et prophanes, divisées en Cing Tomes, le premier
par P. Boaistuau: Tome Premier. Paris, 1582.’ That author
illustrated his article with a figure of the animal, of which a copy
is reproduced in the Cyclopedia. This animal (Broderip writes)
the author states he saw in England in the reign of Elizabeth,
but the probability is that he was deceived by the English bear-
wards and dog-fighters of Hlizabeth’s time, and that some dog
selected for its bear-like appearance in certain points, an
appearance aided by cropping the ears and tail and other skilful
artifices, was palmed upon him and upon others as a hybrid
engendered between a dog and a bear. As Boaistuau’s book is
rare, I have transcribed the passage from the copy in the British
Museum Library (ed. 1566, ch. xxix.) :—
“Gest animal monstrueux, que tu vois figuré au commencement
de ce chapitre, est engendré d’un Dogue d’Angleterre et d’un
Ours: de sorte qu il participe de Pune et de l’autre nature: Ce
qui ne semblera estrange a ceux qui ont observé a Londres
comme les Dogues et les Ours sont logez en de petits cachots, les
uns aupres des autres: et quand ils sont en leurs chaleurs, ceux qui
sont deputez pour les gouverner, enferment une Ourse et un Dogue
ensemble, de sorte que pressez de leurs fureurs naturelles, ils
convertissent leur cruauté en amour, et de telles conjonctions
nayssent quelquefois des animaux semblables a cestuy, encore que
1907. | MR. H. SCHERREN ON HYBRID BEARS. 433
soit bien rarement: entre lesquels en ay observé deux, quon
avoit donné 4 monseigneur Je marquis de Trans: l'un desquels il
fist present 4 monsieur le Conte d’Alphestan, ambassadeur de
VEmpereur : l'autre quwila faict amener en France, sur lequel i’ay
se retirer cestuy au naturel, sans que Je peintre y ait rien
obmis.”
The first well-established case of hybridity in Bears appears to
have occurred in the Society’s Gardens *, between a Black Bear ¢
(Ursus americanus) and a European Brown Bear ¢ (U. arctos).
Copulation was observed in May 1859, and on December 31st
three cubs were born, “ naked and blind, and about the size of a
full-grown rat.” One cub was carried by the dam in her mouth
for a day or two; and as it disappeared it was supposed that she
devoured it. At the age of five weeks the surviving cubs (¢, 2 )
were ‘‘as large asa common rabbit. Their eyes began to open
by this time; they were covered with a short thick fur, and were
nearly black.” This last observation is of some value, as it
supports those of other authorities with regard to the coloration
of the hybrid cubs following that of the male. Mr. Bartlett did
not give the duration of life, but reference to the Occurrence
Book shows that they died on Feb. 14, 1860.
The next case occurred in the Zoological Garden, Cologne, the
male parent being a European Brown Bear (U. arctos) and the
female a Grizzly Bear (U. horribilis). Prof. H. Alexander Pagen-
stecher, of Heidelberg, visited the Gardens in Paris, Lyons, and
Cologne, and the following passage f occurs in bis account of what
he saw in the city last-named :—
“Von Raubthieren haben wir namentlich noch den Baren-
zwinger, ausgezeichnet durch die jungen Bastarde vom gewohn-
lichen bene Biren und der grauen Birin, welche in dem
weiten Kifige trotz ihres gewaltigen Korpers mit solecher Schnel-
ligkeit umherrannte, dass man von der eminenten Gefihrlichkeit
des Thieres in der Freiheit eine geniigende Vorstellung zu erlangen
vermochte.”
There seems to be no doubt about the event, but particulars
were not entered at the time in the records of the Garden.
Dr. Wunderlich, in reply to my enquiries, regretted that he could
add nothing to the foregoing account, and informed me that no
one now in the Garden remembered the hybrids.
Hanover Zoological Garden is said to have been the scene
of the third case, about which, however, some doubt exists.
Dr. Theodore Kottnerus-Meyer, writing on Mammalian Hybrids §,
says :—
“In den achtziger Jahren .... besass der Hannoversche
* A.D. Bartlett, P. Z. S. 1860, p. 130.
+ * Zoologischer Garten,’ 1867, pp. 287, 288.
t * Zoologischer Garten,’ 1904, p. 61.
434 MR. H. SCHERREN ON HYBRID BEARS. [ Apr. 23,
Garten auch eine Grislybiirin (UV. ferox) die in gliicklichster Ehe
mit einem Braunbiiren (U. arctos) lebte, und wiederholt von ihm
mit Erfolg gedeckt wurde. Die Jungen glichen dem Vater und
hatten ganz den Typus des Braunbiren.”
Dr. Schiff, who has been Director since 1893, can find no trace
of these hybrids, but has kindly promised to make enquiries.
The hybrids now in the Society’s Gardens were born in the
Garden which formerly belonged to Herr Nill, who bred hybrids
between Polar and Brown Bears since 1874. His original stock
consisted of a Polar Bear ¢, about 33 years old, and a Brown
Bear 2, a year younger. Having shown a friendly disposition
to each other, playing as freely as they could through the bars,
the experiment was made of putting them tugether, with the best
results. In the summer of 1875 copulation was observed, and on
January 9, 1876, two cubs were thrown, which were quite white.
The colour, however, soon changed to a silver grey with a bluish
tinge, and by the summer to dark brown with a similar tinge.
There was no sign of the whitish neck- or nape-band, generally
more or less noticeable in young Brown Bears; and by the end
of the summer the coats were yellowish white. Two other cubs
were born from the same parents on January 14, 1877, and the
following report on them by Dr. Steudel and Herr E. v. Martens *
is worth quoting in full :-—
“Die beiden halbjihrigen sind gegenwartig vorherrschend
graubraun, doch etwas ungleichmiissig, die Kehlgegend in ihrer
ganzen Ausdehnung auffallig hell fast weisslich. Die beiden
anderthalbjihrigen sind viel heller, Riicken und Seiten isabell-
farbig, ein dunkelbrauner Mittelstreifen, bei dem einen ziemlich
breit tiber den ganzen Riicken sich erstreckend bei dem andern
nur in vorderen Theil schwach angedeutet, Oberseite des Kopfes
hellbraun, Unterseite des Kopfes und Rumpfes weisslich, alle vier
Extremititen noch ziemlich dunkelbraun.”
Two other lots of cubs were born from the same parents; and
then one of the female hybrids was paired with the Polar Bear
(her sire), and produced in succession several litters of three-
quarter blood Polar Bears. The old female Brown Bear was
given a mate of her own species, and produced normal cubs. It
was established that the half-blood Polars were fertile inter se, as
were the females with a pure Brown male.
The original Polar Bear died in the summer of 1898, and the
female Brown Bear, which had thrown more than fifty cubs, was
shot when the Garden was closed in 1906. At that time the
hybrids (Polar and Brown) now in the Society’s Gardens were
purchased. Both are females, one half- and the other three-
quarter Polar. The half-blood was born in December 1893, and
* © Zoologischer Garten,’ 1877, p. 402.
1907.4 MR. H. SCHERREN ON HYBRID BEARS. 435
in December 1897 she threw the three-quarter blood cub to her
own sire, the original Polar Bear, which it may be mentioned was
obtained by von Heuglin’s expedition.
In his last Guide (undated) to his little Garden, Herr Nill thus
described the animals :—
“Wiahrend die Bastardmutter gelblichbraun mit helleren
Abzeichen am Kopf und dunkleren auf dem Riicken und an den
Fiissen gefirbt ist, in den Korperformen aber noch zwischen His-
und Braunbiir steht, hat das dreiviertelbliitige junge Tier voll-
standig die Gestalt und die Farbe des Hisbiren angenommen
und ist nur noch durch eine ganz hellbraune Schattierung lings
des Riickens von einem solchen zu unterscheiden.”
Since then the young has grown darker, but were it possible to
put her toa Polar Bear her cubs would probably be indistin-
guishable from those of the true Ursus maritimus. When I
visited the pretty Garden at Halle-an-der-Saale last summer, the
Director, Dr. Gustav Brandes, drew my attention to a cage con-
taining a male Polar Bear and a female Brown Bear, and remarked
that he had reason to believe they had produced young, which
had been eaten by the dam. MHarly in February of this year he
kindly informed me that cubs had been born, and the mother was.
caring for them. The animals were put together in 1902;
copulation was observed in the summer of 1904, and at the close
of the year the female withdrew into the inner compartment,
remaining there for some days, but 1t was impossible to be certain
that a birth had taken place, though the condition of the mamme
rendered that probable. Pairing again took place in the summer
of 1905, and there was the same uncertainty as to results.
Dr. Brandes, however, assumes that in both cases cubs were born
and eaten by the dam, whose instinct of fostering her young had
not developed. In July 1906 pairing was observed, and on
January 23, 1907, three white cubs were born. Lest I should
unintentionally misrepresent Dr. Brandes’ view I quote a passage
from his letter textually :—‘‘ Mich freut es, dass ich wieder
einmal meine Ansicht, dass sich das Brutpflegeinstinkt erst
entwickeln muss, mal wieder gliinzend bestiitigt hat. Man kann
sich keine bessere Mutter denken, und daher hat sie doch
zweimal die Jungen gefressen.” By theend of February the dark
dorsal stripe was present. These cubs will be kept under close
observation, and photographed from time to time, in order to
illustrate the colour-changes in the coat. For more than a year
Dr. Brandes has kept a Korean female Black Bear (Ursus tor-
quatus) with a pair of Sloth Bears (Jelursus ursinus), and they
have lived peaceably together, though no young have been
produced.
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Acrobates, 233.
Acrodytes
daudinii, 328.
Acryllium
vulturinwin, 170.
Addax :
naso-maculatus, 2.
Aipyornis, 231, 234.
Adpyprymnus
rufescens, 202,
217, 222.
/Krocharis, 360.
Agathodes
ostensalis, 82.
Agrotera
eudoxantha, 78.
flavibasalis, 78.
Jumosa, 78.
pictalis, 78.
semipictalis, 78.
Akodon
caliginosus, 142.
Alactaga, 232.
Alcelaphus
bubalinus, 2.
caama, 114.
Alestes
imbert, 307.
Alma, 415.
Amara, 323.
Amblysomus
chrysillus, 288.
obtusirostris, 288.
Amphibolurus
barbatus, 58.
Anguilla
bengatlensis, 311.
labiata, 311.
Anguis, 57.
Anomalurus, 228, 229.
Antilocapra
americana, 114.
Arachnidium, 252, 255.
203,
205, 206, 207, 208,
INDEX.
Arachnoidia, 252.
ray-lankesteri,
251, 255, 257.
Archeopteryx, 232, 233,
234, 235, 236.
Archernis
callixantha, 86.
eucosma, 86.
ignealis, 86.
Arctogalidia
trivirgata, 198.
Arctomys
marmotta, 212.
Artamia, 353, 354, 357,
358, 359, 305, 366,
371.
Artamus, 353,
359, 865.
leucogaster, 362.
Arvicanthis
dorsalis, 294.
Arvicola
rutilus, 413.
Atherura
africana, 210.
Aulacodes
acroperalis, 76.
bipunctalis, 76, 87.
diopsalis, 76.
gontophoralis, 76.
lunalis, 76, 87.
plicatalis, 76.
purpurealis, 76, 87.
trichoceralis, 76.
Aulacorhamphus
sulcatus, 1.
250,
307, 358,
Barbus
bructi, 309, 311.
elephantis, 310.
eutenia, 308.
holubi, 309.
inermis, 310.
kessleri, 308.
Barbus
natalensis, 309.
polylepis, 308.
rapax, 507.
sector, 309, 311.
trimaculatus, 310.
Barentsia
misakiensis, 252.
Benhamia, 415.
Bettongia
ogilbyi, 205, 207.
penicillata, 205, 207.
Bitis
arietans, 51.
gabonica, 51.
nasicornis, O1.
Bocchoris
adipalis, 79.
aurotinctalis, 79.
euphranoralis, 79.
invertalis, 79.
telphusalis, 79.
Beotarcha
hyalinalis, 86.
limbata, 86.
Bombinator, 327.
ortentalis, 414.
Bos, 116.
grunniens, 304.
Botyodes
asialis, 81.
flavibasalis, 81.
Bradina
glaucinalis, 77.
ampresalis, 77.
melanoperas, 77.
Bramatherium, 114.
Bubalis
caama, 184.
Buchanga, 367.
Bufo, 335, 337.
agua, 336, 337, 348.
marinus, 336.
vulgaris, 414,
438
Bullimus
bagobus, 141.
Buteo
desertorum, 171.
Calamachrous
albipunctalis, 86, 87.
tranquilalis, 86.
Calamites
cyanea, 328.
Calliste
Fastuosa, 143.
Calogale
rutilus, 6.
Calyptorhynchus
banksi, 380.
Campophaga, 361.
Campylognathus, 225
Canis
anthus, 173.
curvipalatus, 403.
Capoéta, 309, 310.
Capra
megaceros, 188.
Caprinia
castanealis, 80, 87.
conglobatalis, 80.
Selderi, 80.
ocellalis, 80, 87.
Capromys
pilorides, 208, 216.
Cephalophus, 115.
coronatus, 114.
grimmi, 185, 298.
maxwelli, 18d.
natalensis, 298.
Ceratophryne
nasuta, 327.
Ceratophrys, 351, 352.
dorsata, 331.
nasuta, 324,
ornata, 336, 347, 348.
Cercoleptes
caudivolvulus, 198.
Cercopithecus
albigularis, 286.
campbelli, 2
denti, 2
mona, 2.
pygerythrus erythrar-
chus, 286.
Cervicapra
arundinum, 237, 299.
— occidentalis, 237.
bohor, 185, 186.
Cervus
aristotelis, 192, 193.
cashmiriensis, 1.
elaphus, 2, 114.
maral, 1.
sika, 192, 216, 219.
INDEX.
Chalcides, 58, 59.
ocellatus, 59.
Chalcidoptera
emissalis, 79.
Chamzleolis,
53.
Chameleon, 58, 67.
basiliscus, 36, 39, 40,
44.
calcarifer, 36, 37, 39,
41, 43, 44.
dilepis, 36, 37, 39, 40,
41, 43, 44.
monachus, 36, 37.
parvilobus, 36, 37, 38,
39, 40, 41, 44.
pumnilus, 36, 39, 41, 42,
43, 44, 45, 68.
teniobronchus, 36, 41,
42, 48, 44, 68.
verrucosus, 36, 37, 38,
40, 42, 44, 49.
vulgaris, SON Ole tle
49,
45-49,
Chlamydopborus, 198.
Chlamydosaurus, 52.
Cirrhochrista
etherialis, 69.
caconalis, 69.
Jiguratalis, 69.
pulchellalis, 69.
Clarias
gariepinus, 311.
Cnaphalocrocis
medinalis, 79.
Cobus
lechwe, 237
robertsi, 237.
smithemannt, 237.
Ceelogenys, 193.
paca, 209, 216.
Coluber
corais coupert, 284.
Compsognathus, 231.
Connochetes
gnu, 184, 190.
taurinus, 184.
Contopus, 354, 358.
Coptobasis
mesopsectralis, 77.
Corvinella, 353, 358,
359, 360.
Cotachena
aluensis, 'T4.
trivitralis, 74.
Craseomys
bedfordie, 405, 413.
Cricetomys
gambianus, 295, 296.
— adventor, 295.
— viator, 295, 296.
Crocidura
jflavescens
290.
sylvia, 290.
Crossarchus
Jasciatus, 291, 292.
— macrurus, 291.
flavidula,
— senescens, 291, 292.
CSTE UR, 198, 199,
Cr unomys
fallax, 142.
‘melanius, 141, 142.
Cryptoprocta
JSerox, 198.
Cuon
alpinus, 2.
Cyclorhis, 375, 377.
Cylindreeeium, 252, 256.
Cymoriza
ustalis, 76.
Cynictis
levaillanti, 198.
Cynomys
ludovicianus, 212.
Cynopterus
sphinx, 8.
Dafila
spinicauda, 173.
Dasypus, 192.
minutus, 198.
villosus, 198.
Dasyurus
mauget, 206, 208.
viverrinus, 206, 207.
Deba
althealis, 81.
surrectalis, 81.
Dendrogale
Srenata, 8.
Dendrolagus
bennettiz, 202, 208.
Dichocrocis
bimacuéalis, 81, 87.
evaralis, 81.
nigrilinealis, 81.
pardalis, 81, 87.
punctiferalis, 81.
xanthocymna, 81.
Dicrurus, 353.
Didelphys, 183, 216,
220.
marsupialis, 200, 216.
nudicaudata, 208.
Dilocareinus, 275.
Dimorphodon, 224, 22
228, 233.
Diplodocus, 231.
Dipus, 219, 232.
Dolichotis,
210.
patachonica, 209, 210,
208, 209,
Doreatherium
aquaticum, 190.
Dracznura
albonigralis, 77.
horochroa, 77.
torridalis, 77, 87.
unicoloralis, 77.
Draco
volans, 229, 230.
Dromeus
nove-hollandie, 172.
Dromicia
nana, 206, 208.
Dryoscopus, 360.
Dytiscus, 323.
Echidna, 219.
Edolius, 366, 367.
forficatus, 365.
EHlaphurus
davidianus, 2.
Elephas, 192.
africanus, 243, 247,
381, 403.
— albertensis, 384, 402,
403.
— capensis, 384, 385,
386, 387.
INDEX.
Hlephas
melitensis, 239, 240,
241, 249, 243,
249.
meridionalis, 239, 243,
249.
mnaidriensis, 246, 247,
249,
namadicus, 247.
planifrons, 403.
priscus, 239, 243.
Eminoscolex, 415, 423.
barnimi, 431.
neumannt, 431.
ruwenzorit, 428, 430,
431.
sylvestris, 451.
toreutus, 431.
viridescens, 431.
Endotricha
albicilla, 74.
persicopa, T4.
Equus, 220.
chapman, 198.
Erethizon
dorsatum, 211.
Erimetopus, 268.
Erinaceus, 220.
algirus, 196, 197.
europeus, 197.
| Eristalis
— cavendishi, 394, 395. |
— cyclotis, 384, 389, |
390, 399, 401.
— knochenhaueri,. 384,
388, 391, 392, 393, |
394, 395, 398, 403. |
— orleansi, 398.
394, 395, 397, 398,
399, 401, 402.
— peeli, 393, 394.
— pumilio, 248, 384,
400.
— rothschildi, 399.
—- selousi, 387, 388,
389.
— ftoxotis, 385, 386,
388, 389, 390.
— typicus, 384, 387.
antiquus, 239, 248,
244, 245, 246, 247,
248, 249, 250.
capensis, d84.
creticus, 238, 239, 240,
241, 248, 244, 246,
249, 250.
cypriotes, 240, 241,
242, 248, 249.
maximus, 239.
tenax, 319.
Eschatocephalus
vespertilionis, 328.
Etiella
Juscalis, 69, 87.
Eumeces, 56, 58, 59.
algeriensis, 5Y.
| Euryceros, 353, 357, 360, |
— oxyotis, 384, 391, |
364, 368, 571.
prevostit, 369.
Eurypyga
helias, 2.
Eutephria
cribrata, 78.
Evotomys, 404.
amurensis, 405, 413.
mikado, 405, 413.
Felis, 145-168.
sp., 6.
angorensis, 150, 151,
167
aurata, 145.
aureus, 151.
badia, 145, 164.
bengalensis, 6, 165.
cerulea, 150.
caffra, 159.
caligata, 159.
439
Felis
catus, 143, 149,
152, 154, 155,
Ge Gre
— ferus, 150.
caudata, 162, 165.
chaus, 162, 164,
166.
domestica,
167.
Ferus, 150.
hispanica, 149, 150.
huttoni, 151.
inconspicud,
155.
Jaguarondi, 145.
longiceps, 151.
lybica, 158, 166.
madagascariensis, 151.
manul, 155, 157, 161,
299, 300, 801, 3802,
304, 305, 306.
nigripectus, 306.
ocreata, 148, 152, 155,
156, 157, 158, 159,
160, 161, 162, 168,
165, 166, 167, 290,
301, 3804, 305, 306.
— agria, 151.
— caffra, 166, 168.
— sarda, 158.
ornata, 165.
pardalis, 172.
pardus, 198.
ruber, 190.
rubiginosa, 165.
sinensis, 150.
striata, 151, 167.
sylvestris, 148, 149, 150,
152, 155, 1o7, 158,
159, 160, 161, 162,
168, 164, 165, 166,
167, 168, 804, 305,
306.
temmincki, 145.
tigris, 157.
torquata, 148, 151,
152, 155, 159, 160,
161, 163, 164, 165,
167, 168.
vulgaris, 149.
150,
160,
165,
145, 150,
151,
Filodes
zanthalis, 80.
Fredericella
cunningtoni, 250, 251,
254, 257.
duplessisi, 254.
pulcherrima, 254.
regina, 254.
suttana, 251, 254.
walkotti, 254.
440
Funambulus
berdmorei, 10.
rufigenis fuscus, 10.
— typicus, 10.
Funisciurus
cepapi, 292.
Sreret, 292.
palliatus, 292, 293.
— ornatus, 293.
— suahelicus, 294.
sponsus, 292, 293.
Galago
conspicillatus, 287.
crassicaudata, 197,
granti, 287.
moholi, 286, 287.
mosambicus, 287.
senegalensis, 287.
Galeopithecus, 227, 228,
229,
Galictis
barbara, 198.
Gallinago
equatorialis, 29, 35.
aucklandica, 28.
australis, 26, 28, 34.
celestis, 12-35.
delicata, 22, 24, 26.
Jrenata, 26, 28, 34.
gallinula, 22, 29, 30.
major, 27, 30.
megala, 31, 32, 33, 34.
nigripennis, 29.
nobilis, 26, 27, 34.
paraguaye, 26, 28.
raddii, 21, 24.
solitaria, 30, 31, 32,
34,
stenura, 25, 27, 33.
Gardullaria, 423.
Gastrosteus
steindachneri, 414.
Gazella
euchore, 187, 188.
Gecko
mauritanicus, 52.
Genetta
sp., 291.
Selina, 199.
letabe, 291.
Geothelphusa, 274,
Geotrupes
spimger, 313, 318.
stercorarius, 318.
typheus, 319.
Gerrhonotus, 55-59.
ceruleus, 56, 57.
Gerrhosaurus, 56.
INDEX.
Giraffa, 100-125.
camelopardalis antt-
quorum, 105, 120.
— cottoni, 121.
—wardi, 105, 119,
122.
reticulatus, 124, 125.
Glauconycteris
papilio, 288.
variegatus, 288.
Glyphodes
actorionalis, 82.
egealis, 82.
agathalis, 82.
amphitritalis, 82.
bicolor, 82.
brunneomarginalis, 84,
87.
cesalis, 82.
canthusalis, 82.
celsalis, 82.
deliciosa, 82.
doleschali, 82.
eurygania, 82.
eurytusalis, 82.
exaula, 82.
exquisitalis, 84, 87.
flavizonalis, 82.
glauculalis, 82.
hilaralis, 82.
hypomelas, 82.
indica, 82.
itysalis, 82.
laceritalis, 84, 87.
laticostalis, 82.
margaritaria, 82.
marinata, 82.
multilinealis, 83, 87.
ophiceralis, 82.
paucilinealis, 83, 87.
perspicualis, 83, 87.
pfeiffere, 82.
pudicasis, 83, 87.
quadristigmalis, 88,
87.
sectinotalis, 82.
stolalis, 82.
subacusalis, 82.
suralis, 82.
tricoloralis, 82.
wmobria, 82.
unionalis, 82.
zelimalis, 82.
Goniorhynchus
gratalis, 81.
Gonoptera
libatrix, 319, 322.
Gorilla
gorilla diehli, 237.
Graucalus, 358, 357,
376.
Gulo
luscus, 198.
Gymnorhina, 353, 356,
357, 359, 360, 361,
365, 367, 368, 370,
371, 372.
organicum, 395, 363.
Heemogregarina
mansoni, 284.
pocecki, 283.
rarefaciens, 284.
refringens, 234,
shattocki, 284.
Halmaturus
bennettii, 202, 203.
Hapale
devillei, 99.
illigeri, 99.
jacchus, 98.
Haplochromis
moffatti, 311.
Hatteria, 59.
Helictis .
personata, 198.
pierrei, 7.
Helix
pellita, 244.
Helladotherium, 118.
Heloderma, 59-68.
horridum, 61.
suspectum, 52, 61, 62.
Hemitragus
jemlaicus, 188.
Heortia
dominalis, 85.
witessoides, 85.
Herpestes
birmanicus, 6.
exilis, 6.
griseus, 198.
javanicus, 6.
pulverulentus, 198.
rutilus, 6.
Hesperornis, 229, 234.
Heterocnephes
scapulalis, '78.
Hippopotamus
amphibius, 1, 380.
liberiensis, 248.
Hipposiderus, 230.
gigas, 230.
Homa, 197.
Hyena
crocuta, 198, 199.
Hydaspitherium, 114.
Hydrocherus
capybara, 212.
Hydrocyon
lineatus, 307.
Hydropotes, 219.
Hydrothelphusa,
275.
Hyla
cerulea, 328.
goeldit, 139.
palmata, 138.
resinifictrix, 185-140.
taurina, 135.
tuberculosa, 135.
venulosa, 139,
1339).
Hylobates
agilis, 143.
Hymenochirus, 327.
Hypsilophodon, 231.
Hypsipyla
Doan 69.
Hypsopygia
postflava, 79.
Hyrax, 219, 222.
capensis, 193,
195.
Hystrix, 208.
cristata, 210, 211, 216,
220.
grote, 3.
hodgsoni, 11.
subcristata, 11.
yunnanensis, 3, Wil
268,
137,
194,
Ichthyornis, 234.
Ictonyx
capensis, 292.
Tguana, 45, 46, 56, 67.
tuberculata, 57, 58, 62.
Tguanodon, 232.
Isocentris
minialis, 86.
miniosalis, 86.
Jaculus
orientalis, 208, 212.
Lacerta, 53, 54, 55, 68.
agilis, 53.
vivipara, 414.
Lachesis
lanceolatus, 50, 51.
Lagomys
roylet, 212.
Lalage. 353, 354.
Taniarius, 354, 357, 360,
361.
poliocephalus, 355.
Lanius, 353, 360.
collurio, 368.
excubitor, 358.
Leggada
minutoides, 295.
INDEX.
Lemur, 220.
albifrons, 197.
catta, VOT.
coronatus, 197.
macaco, 197, 209.
mongoz, 197.
varius, 197.
canthomystax, 197.
Leptobrachium, 325, 330.
Leptodactylus, 337.
hexadactylus, 337.
Lepus
capensis, 297.
— aquilo, 297.
— centralis, 297.
— granti, 297.
curopeus, 212.
ochropus, 297, 298.
timidus, 405, 414.
— ainu, 405.
wassali, 11.
| Lepyrodes
perspicata, 85.
| Leucinodes
apicalis, 85.
Limnias
ceratophylli, 251.
Limnomys, 141.
Limnothelphusa, 266,
267.
maculata, 267.
Liolemus, 67.
magellanicus, dT.
Loecastra
castanealis, 72, 87.
Lophopodella
thomasi, 251, 257.
(Pectinatella) carter?, |
251.
Loris
gracilis, 4.
Lucilia, 323.
Lutra, 220, 286.
vulgaris, 198, 199.
Lycaon
pictus, 2.
| Maealia
apicalis, 71, 87.
caradriniformis, T0, 87.
curtulalis. 70, 87.
margarita, 10.
nyctichroalis, 70.
enochroa, 70.
perdentalis, 71, 87.
picta, 70.
pomalis, 70, 87.
porphyrealis, 71, 87.
seminivea, 70.
tenebrosalis, 71.
unipunctalis, 70, 87.
441
Macropus, 205, 207, 218,
220.
agilis, 201, 207.
alligatoris, 202, 207.
antilopinus, 202, 207.
bennettti, 207.
brunt, 201, 207.
derbianus, 202, 207,
216.
dorsalis, 201, 202, 207,
209.
giganteus, 202, 208,
20%
melanops, 201, 207.
melanotis, 208.
rufus, 202, 207.
ualabatus, 201,
207.
Macroscincus, 58, 59.
Macroxus
annulatus, 298.
leucopus, 8.
Manis
javanica, 11.
Marasmia
Fusifascialis, 79.
Maruca
testulalis, 86.
Megalophrys, 324, 335,
308, 339, 340, 343,
344, 345, 346, 347,
349.
chysti, 325.
fe@, 325, 330.
longipes, 326, 327, 828.
montana, 325, 326, 327,
28, 829, 349.
nasuta, 325, 326,
328, 329, 330, 331,
332, 340, 341,
344, 346, 348, :
351, 352.
Melanomys, 142.
Mellivora
ratel, 292.
Melolontha, 322, 323.
vulgaris, 318.
ZO
| Melursus
ursinus, 435.
Meroctena
staintoni, 8d.
Meta
menardi, 319.
Metrea
nebulalis, 85.
Micralestes
acutidens, 307.
Microchetus
colletti, 277, 278, 279.
griseus, 277, 281.
nucrochetus, 278.
449
Microchetus
culuensis, 279, 280,
281.
Micromys
agrartus, 412.
geisha, 412.
— celatus, 412.
— geisha, 404, 412.
— hokkaidi, 405,
412.
— yakur, 412.
spectosus, 404, 412.
— ainu, 408, 412.
— giliacus, 405, 411,
412.
— navigator, 412.
— peninsularis, 411,
412.
— speciosus, 412.
Midas
devillei, 99.
elegantulus, 91.
erythrogaster, 91.
fuscicollis, 88, 89, 96.
griscovertex, 88, 89, 90, |
91, 92, 93, 98.
illigeri, 88, 93, 99.
imperator, 88, 93, 94.
labiatus, 88, 91.
mystax, 88, 98.
piuleatus, 88, 97, 98.
—- juruanus, 98.
rufiventer, 88, 89, 90,
91, 92, 93.
thomas, 89, 92.
weddellii, 88, 93.
Monodonta, gen. nov.,
passalis, 69, 87.
Moschus, 219.
moschiferus, 190.
Mungos
albicaudis, 291.
cafer, 291.
gracilis, 236.
(Ichneumia) grandis,
291.
Mus
sp., L41.
albigularis, 141.
bagobus, 141.
chrysophilus, 295.
coucha,. 295.
norvegicus, 404, 405,
411.
rattus, 141, 295.
sylvaticus, 412.
vuleani, 141.
Muscicapa, 356, 360.
Myiarchus, 366.
arundinicola, 354.
| Myotis
| Myoxus
| Nosophora
| Nycteribia
INDEX.
Myopotamus
coypu, 181, 212, 213,
214, 215, 216, 217,
223.
mystacinus, 314, 404,
407.
lineatus, 411.
Myrmecophaga |
Jubata, 198.
Nacoleia
charesalis, 81.
Nandinia
binetata, 198.
Nasua
rufa, 198.
Nebria
brevicollis, 318.
Nemorhedus
caudatus, 281.
goral, 114.
swettenhami, 188.
Nesotragus
livingstonianus, 299.
zuluensis, 299.
Neumanniella, 415.
ruwenzorit, 424, 426,
427.
siphonocheta, 424,
425.
tenuis, 424, 425.
Nevrina
procopia, 80.
Newtonia, 360, 365, |
371. |
brunneicauda, 369. |
barbata, 79.
dispilalis, 79.
fulvalis, 79.
semitritalis, 79.
hermannt, 323.
Nycticebus
cinereus, 5.
coucang, 4.
— cinereus, 4.
pugmeus, 4, 5, 11.
tardigradus, 4, 197.
Nyctodactylus, 226, 2277,
228. |
Nyctosaurus, 226, 228.
Okapia, 100-115,
135.
johnston, 128, 129.
126- |
liebrechtsi, 129.
Omphisa
ingens, 87.
Onychogale
Frenata, 206, 208.
Ophiosaurus, 56, 57.
apus, D7.
Opisthocomus, 235.
Oreotragus
saltator, 114.
Ornithocheirus, 227,
229.
Ornitholestes, 231, 235.
Ornithomimus, 232.
Orphnophanes
euceralis, 77.
Orthaga
chionalis, 73, 87.
columbalis, 74, 87.
Juscofascialis, '74, 87.
polychroalis, 73, 87.
seminivea, V4.
vitialis, 73.
Oryx
beatrix, 185, 221.
leucoryx, 185.
Otocolobus
manul, 802.
mongoticus, 302.
— nigripectus, 302.
nigripectus, 302.
Otocyon, 408.
Otomys
irroratus, 294.
— cupreus, 294.
Ourebia
nigricaudata, 185.
Ovibos
moschatus, 114.
Ovis
cowant, 238.
stonei, 238.
tragelaphus, 114, 220.
Pachynoa
spilosomoides, 86.
thoosalis, 86.
Pachyzancla
desmioides, &6.
pallidalis, 86.
Pagyda
paraphragma, 78.
Palzornis
sp., 170.
Paludicella, 252.
erecta, 256, 257.
Paludicola
bibronti, 328.
Papio
maimon, 1.
porcarius, 286.
Paradisea, 353, 354, 356,
359, 371, 372.
raggiand, 363.
Paradoxurus, 4.
minor, 6.
Paramelania, 250, 255.
Parascolex, 426.
Parathelphusa, 275
Parthenodes
rectangulalis, 77, 87.
Pedetes
cafer. 282, 296.
Pelobates, 326, 327, 333. |
335, 337, 340,
342, 344, 350,
341,
351,
cultré ipes, 3ol.
fuscus, 326, 329,
347, 349.
Pelodytes, 338, 340,
342
punctatus, 339, 341.
Perameles
obesula, 206, 208.
Perodicticus
potto, 197.
Petaurista
lylei, 3, 8.
yunnanensis, 5, 8.
Petaurus, 228.
breviceps, 206, 208.
Petrodromus, 285.
schwanni, 289, 290.
sultan, 289, 290.
Petrogale
penicillata, 205, 207.
xanthopus, 205.
Phacocherus
ethtopicus, 188, 189.
Phascologale
penicillata, 206, 208.
Phascolomys
mitchelli, 204, 205,
207.
Phelsuma, 58.
Pheretima, 424.
Philohela
minor, ot.
Phryganodes
basalticalis, 80.
glyphodalis. 80.
maculicostalis, 80.
margarita, 81.
obscurata, 80.
odontosticta, 80.
Phrynosoma, 46.
Phycita
dinawa, 69, 87.
Physignathus, 52.
Phytotoma, 3538, 354,
356, 357.
334, |
341,
INDEX.
Piezorhynchus, 360.
Piletocera
inconspicualis, 78, 87.
Pilocrocis
anigrusalis, 79.
coptobasis, 79.
Pionea
ablactalis, 86.
albifimbrialis, 86.
bractealis, 86, 87.
nypsiusalis, 86.
Pipa, 327, 332, 333, 342,
343, 344, 345, 352.
Pipistr ellus
NANUS, 288.
pipistrellus, 318.
Pitangus, 353, 357, 365.
Pitta, 353, 367.
baudi, 362.
Pityriasis, 353, 360, 361,
365, 367, 368, 371.
Platythelphusa, 266, 267,
268, 274, O75.
armata, 259, 266, 267,
268, 269, 271
274, 276.
conculcata, 259, 268,
, 272,
273, 276.
maculata, zon 268,
270; Ql, 273; 74,
275, 276.
Plecotus
auritus, 405, 406.
| Pleuronectes
jflesus, 175, 180.
macrocephalus, 180.
platessa, 180.
Plumatella
repens, 250, 251, 253,
tanganyike, 250, 251,
252, 257.
Polychrus, 46.
marmoratus, £7.
| Polygrammodes
grossalis, 87.
purpuralis, 87.
Polylophota
barbarossa, 72.
truncalis, 72, 87.
Polythlipta
divaricata, 85.
macralis, 85.
| Polytoreutus, 429.
bettonianus, 416.
ceruleus, 422.
granti, 420, 421, 422,
493.
gregorianus, 420.
kirimaensis, 415, 417,
418, 419, 420, 422,
423.
443
| Polytoreutus
magilensis, 422.
ruwenzorit, 415, 416,
417, 418, 419, 420,
421, 422, 423.
sylvestris, 415, 416,
417, 418, 419, 422,
423.
usindjaensis, 415, 418,
Porcula
salvania, 190.
Potamocarcinus, 275.
Potamocherus
africanus, 190.
cheropotamus nyase,
298.
Potamon,
275.
ambiguus, 265.
hilgendorfi, 261, 263.
johnstoni, 260, 261,
263.
mrogoroensis, 265, 266.
perlatus, 268, 264.
suprasulcatus, 265.
(Geothelphusa) emini,
274.
267, 268, 274,
(Parathelphusa) 2élo-
ticus, 274, 276.
(Potomonautes), sp.,
259, 262, 266.
(—) celebensis, 262
2638.
—) inflatus, 259.
(—) orbitospinus, 259,
260, 261, 274, 276.
(—) perlatus, 268, 264
276.
(—) platynotus, 259,
264, 265, 266, 270
276.
Potamonautes, 268, 274.
Pottsiella, 252, 256.
Presbytes
sp., 4.
nigripes, 4.
Prionops, 353, 354, 357,
359.
Prozwdema
inscisalis, 86.
Procyon
cancrivorus, 198.
lotor, 198, 199.
Proteles
cristata, 198.
Protoceros, 114.
Pseudaspis
cana, 284.
Pseudochirus
sp., 206, 208.
444
Pseudochirus
_peregrinus, 206, 220.
Pterodactylus, 225, 226,
228.
Pteromys,
229.
russicus, 409.
yunnanensis, 8.
Pteropus, 226, 229.
vulgaris, 229.
Pterostichus
niger, 319.
Ptilocerus
lowi, 233.
Pycnarmon
jaguaralis, 78.
lactiferalis, 78.
Pygospila
bivitralis, 85.
imperialis, 85, 87.
marginalis, 85, 87.
Pyrausta
ceadesalis, 87.
celatalis, 87.
eriopisalis, 87.
triticalis, 87.
Python
molurus, 283.
spilotes, 284.
227
’ tt}
Rana
amurensis, 414.
esculenta, 328, 331,
332, 339, 345, 348.
— chinensis, 414.
guppyi, 331, 333, 384,
337, 338, 3839, 342,
343, 344, 345, 346,
348, 349, 350, 351,
352.
temporaria, 414.
tigrina, 331, 334, 337,
345, 348.
Rehimena
phrynealis, 78.
Rhacophorus
sp., 348, 352.
Rhamphoccelus
brasilius, 148.
Rhamphorhynchus, 225,
228, 229, 2338.
Rhaphicerus, 185, 191.
horstocki, 237.
— natalensis, 237.
melanotis, 184.
newumanni capricornis,
298.
Rbimphalea
heranialis, 79.
linealis, 79, 87.
INDEX.
Rhimphalea
scelatalis, 79.
trogusalis, 80.
Rhinolophus
cornutus, 405, 406.
JSerrum-equinum, 312,
323, 324.
— nippon, 405, 406.
hipposiderus, 312, 314,
323.
Rhizomys
pruinosus, 10.
Rhombus
maximus, 178.
Rhynchocyclus, 354, 358.
Sacada
inordinata, 75.
nigropunctata, 75.
Saccostomus
campestris, 294, 295.
Suscus, 294, 295.
Sameodes
tristalis, 85, 87.
Samotherium, 113.
Sayornis, 358, 364, 866.
cineracea, 355, 362.
Scaphiopus, 327, 541.
solitarius, 328.
Scaphognathus, 229.
Sciuropterus
buechneri, 410.
momonga, 409.
— amygdali, 409.
russicus, 404, 409,
410.
— athene, 404, 409.
Sciurus, 220.
americanus volans, 409.
berdmorei, 10.
bicolor, 212.
griseimanus, 8, 9.
leucopus, 8, 9.
— fumigatus, 9.
— typicus, 9.
nacclellandi, 4.
— barbei, 4.
— maritimus, 4, 9.
rodolphi, 4, 10.
maximus, 212.
mindanensis, 141.
mouhoti, 10.
orientis, 410.
palimarum, 212.
rodolphi, 10.
vassali, 9.
vulgaris, 212.
— ortentis, 405, 410,
411.
— rupestris, 404, 410.
Scolopax
rusticola, 34.
Scotophilus
damarensis, 287.
kuhlii, 8.
minimus, 288.
nigrita, 287, 288.
— herero, 287.
planirostris, 287.
schlieffeni, 288.
viridis, 287.
Scotosia
dubitata, 319, 322.
Semnopithecus
nigripes, 4.
Seps, 58.
Sicista
caudata, 404, 405, 413,
414.
concolor, 413, 414.
leathemi, 413, 414.
subtilis, 414.
tianshanica, 414.
Sivatherium, 113.
Sorex
araneus, 407.
buxtoni, 408.
daphenodon, 404, 407. .
gracillimus, 404.
minutus, 404, 409.
— gracillimus, 408.
pygmeus, 409.
shinto, 408.
— se@vus, 404, 405,
408.
unguiculatus, 404, 405,
407.
Sparganum
baxteri, 282.
hematobium, 283.
japonicum, 283.
mansont, 283.
Sphecotheres, 358, 376.
Spinturnix
vespertilionis, 324.
Spongilla
tanganyike, 250, 256.
Stericta
cornucalis, 73, 87.
divitalis, 72.
flammealis, 72, 87.
harraldusalis, 72.
prasina, 72.
sporeta, 72.
subviridalis, 73, 87.
Streptospondylus, 231.
Suricata, 220.
tetradactyla, 199.
Sus
sp., 142.
scrofa, 190.
Sylepta
acridentalis, 82,
aurantiacalis, 81.
chalybi fascia, 81.
chromalis, 81.
dissipitalis, 81.
multilinealis, 81.
ochrifusalis, 82.
pernitescens, 81.
solilucis, 81.
Syngamia
ampliatalis, 79.
floridalis, 79.
marmorata, 79.
Tachyphonus
coronatus, 143.
Teeniopterus, 358.
Talanga
tolumniats, 77.
Tamandua
tetradactyla, 198.
Tamias
asiaticus, 404, 411.
lineatus, 405, 411.
Taphozous, 225.
Tarentola, 58.
Tatera
bechuane, 294.
lobengule bechuane,
294,
Taurotragus
oryx, 380.
Telendrilus, 423.
Termioptycha
cyanopa, 74.
Terpsiphone, 356, 559,
360, 362, 366.
Tetraceros, 114, 115.
Thamnomys
arborarius, 295.
dolichurus, 295.
Thliptoceras
octoguttale, 85.
Thrasaétus
harpyia, 281.
Thryonomys
swinderenianus, 286.
Thylacinus
cynocephalus, 207.
Tilapia
natalensis, 311.
sparrmant, 311.
Tiliqua, 53, 54, 55, 57.
scincoides, 49, 54, 58,
59.
INDEX.
Tipuliforma, gen. nov., 75.
triangulalis, 75, 87.
Totanus
glareola, 50,
Trachydosaurus, 53, 55,
58.
rugosus, 8.
Tragulus, 193, 195.
affinis, 11.
kanchil affinis, 11.
— pierre, Il.
meminna, 191, 192.
ravus, 11.
Tribelesodon, 227, 228.
Trichelurus, 302.
Trichosurus
vulpecula, 203, 208.
Triphena, 319.
orbona, 322.
pronuba, 322.
Tropidonotus
vibakari, 414.
Tupaia
belangeri, 7.
chinensis, 3, 7.
concolor, 7.
everetti, 140.
JSerruginea, 4.
Frenata, 8.
Tupinambis, 58,
Tyias, 353, 361, 370, 371,
376.
Tyrannus, 353, 361, 366,
370.
Tyspanodes
radiata, 80, 87.
Ulopeza
cruciferalis, 79, 87.
Urogale
cylindrura, 140.
everetti, 140,
Uromastix
acanthinurus, 58.
Ursus
americanus, 433.
arctos, 438, 434.
Serox, 434.
horribilis, 433.
maritimus, 435.
torquatus, 435.
Valdivia, 275.
Varanus, 48, 55, 59-68.
bengalensis, 65.
exanthematicus, 63, 64,
65, 67.
Proc. Zoot. Soc.—1907, No. XXX.
Varanus
gouldit, 63.
griseus, 63, 65, 66.
niloticus, 5d, 63, 64,
65, 67.
ocellatus, 65, 67.
salvator, 63.
Varicorhinus
ansorgit, 311.
bructt, 310, 311.
Vespa, 323.
Vespertilio, 228.
sphing, 8.
Victorella, 251, 253, 256.
pavida, 252, 256, 257.
symbiotica, 251, 259,
2511.
Vipera
berus, 414.
russelli, 51.
Vireo
olivacea, 362,
Vireolanius, 353, 354,
BH), BU lly Boy OUT.
leucotis, 362.
Vitessa
griseata, 75.
suradeva, 75, 87.
zemira, Ta.
Viverra
civetta, 198, 290.
qalaccensis, 6.
megaspila, 6.
Viverricula
malaccensis, 6.
Wellcomia
mitchelli, 282.
Xenophrys, 330.
monticola, 340, 342,
302.
Xenopirostris
pollent, 376.
Xenopus, 327, 344, 345,
346.
Xylophasia
polyodon, 322.
Zamenis
flagelliformis, 284.
Zinekenia
Jascialis, 78.
Zonurus, 52-55, 67, 68.
cordylus, 53.
gigunteus, 48, 49, 52,
IO,
30
P. Z. S. 1907, pp. 1-236,
were published on June 12th, 1907.
Printed by Tayror and Francis, Red Lion Court, Fleet Street.
No. 41.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
March 5th, 1907.
FreDERIcK GiuLetr, Esq., Vice-President, in the Chair.
The Hon. WALTER RoruscuH xp, M.P., F.Z.S.,exhibited amounted
specimen of a Gorilla, Gorilla gorilla diehli.
Mr. RoruscuiD also described the Natal form of the Steinbuck
as being darker and more vinaceous rufous in colour, and the
North Rhodesian Reedbuck as having greyer body and less rufous
head and stouter horns than the typical forms; a new Lechwe
allied to both Cobus lechwe and C. smithemanni, but with black
patches only on the shoulders and black hairs scattered about
the neck; and a new Sheep allied to Ovis stonei, but almost
entirely black.
A communication from Miss Dororuza M. A. Bare contained
an account of the discovery, in cave-deposits in Crete, of remains
of Elephants, some of which were referred to a new species.
Mr. Cuarues F. Rousseter read a report on the Polyzoa of the
Third Tanganyika Expedition. Five species were represented in
the collection, three of which were described as new. Of the
five species, three belonged to the Phylactolemata and two to the
Gymnolemata. Amongst the latter was <Arachnoidia ray-
lankestert Moore, which was found in some abundance on shells
dredged from deep water.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications
along with the ‘Proceedings’; but it may be obtained on the day of publication
at the price of Stxpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance.
14
Dr. W. A. Cunninerton, F.Z.8., read a report on the Brachyurous
Crustacea of the Third Tanganyika Expedition. The collection
contained specimens from both Nyasa and Tanganyika. Including
a few individuals which had hitherto passed without notice in the
collection of the British Museum, there were now on record three
species from Nyasa and five from Tanganyika. Of these species,
three were described as new. The forms from Nyasa all belonged
to the widely distributed subgenus Potamonautes ; but while two
species from Tanganyika also belonged to that subgenus, the lake
contained three species belonging to the remarkable endemic
genus Platythelphusa A. Milne-Kdwards. The suggested marine
appearance of P. armata was considered to be only superficial,
and the peculiar character of the Brachyuran fauna of
Tanganyika could be explained on the grounds of a prolonged
isolation of the lake.
Mr. F. E. Bepparp, F.R.S., described two new species of
African Oligocheete Worms of the genus Microchetus belonging
to the collection of the Christiania Museum.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 19th March, 1907, at half-past Hight
o'clock P.M., when the following communications will be made :-—
1. Dr. L. W. Sampon, F.Z.S.—On some new or little-known
Animal Parasites.
2. Mr. Herpert F. Stanptne, M.Sc.—On recently Discovered
Sub-fossil Prosimiz from Madagascar, their Affinities with extant
Lemurs and with the higher Primates.
3. Messrs. OupDFieLD THomas, F.R.S., F.Z.8., and R. C.
Wroveuton, F.Z.8.—The Rudd Exploration of South Africa.—
VII. List of Mammals obtained by Mr. Grant at Coguno,
Inhambane.
The following Papers have been received :—
1. Mr. C. J. Wira.—An Account of the South American Cheli-
ferine in the Collections of the British and Copenhagen Museums.
2. Mr. T. A. Cowarp, F.Z.S.—On the Winter Habits of the
Greater Horseshoe and other Cave-haunting Bats.
Communications intended for the Scientific Meetings of the
ZooLoGIcAL Society or Lonpon should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 HLANovER Square, London, W.
March \2th, 1907.
No. 42.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
March 19th, 1907.
Dr. Henry Woopwarp, F.R.S., Vice-President, in the Chair.
The Secrerary read a Report on the additions that had been
made to the Society’s Menagerie during the month of February
1907.
Mr. Herpert F. Stanpine read a paper, illustrated by lantern-
slides and large series of photographs and specimens, on recently
discovered subfossil Prosimiz from Madagascar, in which he dis-
cussed their affinities with extant Lemurs and with the higher
Primates. The remains were obtained in the muddy bed of a swamp
formed by the blocking-up of the river Mazy by a lava-flow, at from
a few inches to 3 or 4 feet below the surface. They consisted of
a large number of skulls and limb-bones of Lemurs and Lemur-
like animals. This great amount of material enabled the author
to correborate the view, previously put forward by Dr. Forsyth
Major, that the extinct Lemurs of Madagascar were, in many
respects, intermediate between existing Lemurs and Monkeys,
and to express his belief that the New World Monkeys and the
Lemuride,as well as the Malagasy Indrisine, had a common origin.
He also stated his epinion that, in view of the recent additions to
our knowledge of the Prosimiz and of what the present collection
revealed with regard to their close relationship to the Apes, it
was not possible to separate the Primates, as hitherto, into the
two suborders Lemuroidea and Anthropoidea.
Dr. L. W. Sampon, F.Z.S., read a paper on animal parasites,
and described three new species as follows :—
WELLCOMIA MITCHELLI, gen. et sp. nov.
Habitat. Small intestine of Pedetes caffer.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Stxrpence, or, if desired, sent post-free for the sum of Sir
Shillings per annum, payable in advance.
16
Only females found, 12-15 mm. long and about 1 mm. broad.
Characterized by the presence of a conical ovipositor 2-3 mm.
long, placed ventrally on the anterior third of the body, 2—
3°5 mm. from the cephalic extremity, and by a spirally twisted
tail, 2-3 ym. long, terminating in a fine point. Body semitrans-
parent. Head tapering anteriorly. Mouth trilabiate ; cesophagus
long and terminating in a spherical bulb. Anus open ventrally
at 3-4 mm. from tail-end. Eggs smooth, oblong, asymmetrical,
and measuring 60-65 p by 28-32 yp.
SPARGANUM BAXTERI, sp. n. ?
Habitat. Connective tissue of Man.
Long, flat, unsegmented body, 15 cm. long and 1°5 mm. broad,
with numerous irregular transverse folds and a distinct longitu-
dinal groove on ventral surface. Anterior extremity 2°5 mm.
broad ; head completely invaginated. Posterior extremity 2 mm.
broad, with shallow median slit. Extracted from an abscess on
the thigh of a Masai, British Central Africa.
ScHISTOSOMUM MANSONI, Sp. N.
Habitat. Blood-vessels of Man.
In the Congo Free State, in other parts of Africa, and in the
West Indies there is a form of Bilharziasis clinically and patho-
logically similar to the Asiatic form caused by Schistosomum
japonicum, and unlike the classic Hast African form due to
S. hematobium. The eggs of the species which causes this peculiar
form are never found in the urine, but seem to be eliminated
through the intestine only. They differ from those of S. hema-
tobium in having a broad lateral spine totally different in size,
shape, and position from the small, straight, terminal spine which
characterises the ova of S. hematobium. Hitherto, the laterally
spined ova, usually observed in Egypt in cases of mixed infection,
have been looked upon as having been d’storted whilst passing
through the rectal mucosa. Sir Patrick Manson suggested several
years ago, that the laterally spimed ova found in the feces of
patients, and never in the urine, might represent a new species.
In appreciation of this, one of his many genial intuitions, the new
species is dedicated to him.
Dr. L. W. Sampon also described five new Hemogregarines
discovered by himself and Dr. C. G. SeLigmann in Snakes, as
follows :—
HMOGREGARINA POCOCKI, sp. n.
Habitat. Erythrocytes of Indian Python, Python molurus L.
Club-shaped, 14-16pJlong. Anterior extremity rounded, 3-15 pu
broad. Posterior extremity attenuated and recurved. Cytoplasm
more or less granular. Nucleus median or nearer posterior ex-
tremity, large, oval, and with coarse deeply staining chromatin
7,
granules. Parasite lies parallel or obliquely to long axis of host-
cell, of which it occupies about two-thirds, without causing much
alteration beyond displacement of nucleus.
H£MOGREGARINA SHATTOCKI, sp. n.
Habitat. Erythrocytes of Diamond Snake, Python spilotes
(Lacép.).
Club-shaped. Some forms more slender, 14-15 yp long and 2 u
broad, with both extremities rounded and differing only slightly
in thickness. Other forms more bulky, somewhat similar to those
of Python molurus, but larger, 224 by 4. Nucleus median
and very large, 9 u by 4. Host-cell sometimes slightly distorted,
nucleus pushed to the periphery.
H#MOGREGARINA REFRINGENS, Sp. n.
Habitat. Erythrocytes of Hoary Snake, Pseudaspis cana L.
Crescentic, bean-shaped, and discoidal forms oceur. The slender
crescentic forms have a long, oval, and more or less central nucleus.
The bean-shaped forms measure 10-12 mu in length by 5-6 p in
width, they have a wide central nucleus, and their cytoplasm is
literally crammed with rounded highly refractive granules. Host-
cell unaltered beyond occasional displacement of nucleus.
H2MOGREGARINA MANSONI, sp. n.
Habitat. irythrocytes of Testaceous Snake, Zamenis flagelli-
formis L.
Oval or bean-shaped cyst 12-13 long by 5-6 broad, en-
closing club-shaped parasite doubled up in the form of a letter U
with both branches of equal length and closely applied. Nucleus
median and situated near bend at one pele of cyst. Chromatin
arranged in transverse parallel lines or in concentric circles.
A characteristic feature is the almost constant presence of two
large chromatoid granules usually placed one on each side of
nucleus. Host-cell unaltered, nucleus slightly displaced.
H@MOGREGARINA RAREFACIENS, Sp. n.
Habitat. Erythrocytes and leucocytes of Couper’s Snake,
Coluber corais var. cowpert Holbr.
1. Slender, elongate, cylindrical forms 14 p long by 1:5 p broad,
sometimes presenting a refringent granule or vacuole at each
extremity. Host-cell apparently unaltered or only slightly
hypertrophied.
2. Large bean-forms 12-13 p long by 4-5 w broad, with
cytoplasm more or less granular and occasionally vacuolated.
Nucleus small, round, median, with fine chromatin grains. The
host-cells containing this form measure about four times the
normal size, are entirely dehemoglobinized and greatly attenuated.
Their nucleus is hypertrophied. Sometimes two or even three
parasites may be found in the same host-cell.
18
A paper by Messrs. Otprienp Tuomas, F.R.S., F.Z.S., and
R. C. Wroventon, F.Z.S., was read, giving an account of a
collection of Mammals, the seventh of the series, made by
Mr. C. H. B. Grant at Coguno, Inhambane, and presented to
the National Museum by Mr. C. D. Rudd. The collection
consisted of 212 specimens belonging to 39 species, of which six
were described as new.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 9th April, 1907, at half-past Hight
o'clock p.m., when the following communications will be made :-—
1. Mr. T. A. Cowarp, F.Z.8.—On the Winter Habits of the
Greater Horseshoe and other Cave-haunting Bats.
2. Mr. G. A. Boutencrr, F.R.S.—On a small Collection of
Fishes made in the Hastern Watershed of the Transvaal by
Capt. G. E. Bruce.
3. Mr. W. P. Pycrart, F.Z.8.—Oontributions to the Osteology
of Birds.—Part VIII. Tyranni, Hirundines, Muscicapze, Laniine,
and Gymnorhines.
The following Papers have been received :—
1. Mr. C. J. Wira.—An Account of the South American
Cheliferine in the Collections of the British and Copenhagen
Museums.
2. Mons. 8. A. Bururiin.—On some new or little-known
Siberian Birds.
3. Mr. F. E. Bepparp, F.R.S.—Notes upon the Anatomy of a
Species of Megalophrys, with references to other Genera of
Batrachia.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Socrery or Lonvon should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 HANOVER Square, Lonpon, W.
March 26th, 1907.
No. 43.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
April 9th, 1907.
Dr. Henry Woopwarp, F.R.S., Vice-President,
in the Chair,
Mr. R. I. Pocock, the Superintendent of the Gardens, exhibited
a photograph and the skull of the specimen of Pallas’s Cat (Welis
manul) that had recently died in the Menagerie.
Mr. G. A. Boutenenr, F.R.S., read a paper on a collection of
Fishes made in the Eastern Watershed of the Transvaal by
Capt. G. E. Bruce and presented by him to the British Museum.
The collection contained specimens of 18 species, of which several
had not been previously recorded from the Transvaal and five
were new.
Mr. W. P. Pyorart, F.Z.8., read a paper on the Osteology
of the Oligomyodian and Diacromyodian Passeres. After
referring to his previous contribution (published in the ‘ Pro-
ceedings’) on the Osteology of the KEurylemid and Tracheophone
Passeres, he remarked that there seemed little room for doubt
but that the Diacromyodian and Oligomyodian Passeres must be
regarded as divergent branches of a common stem.
The latter suborder included the Tyranniformes, Phytotomide,
and Pittide, while the former embraced the remaining Passeres.
In the present communication some fourteen Families were
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Sixpence, or, it desired, sent post-free for the sum of Sie
Shillings per annum, payable in adyance. 3
20
described, and these were divided into four groups—Hirundines,
Muscicapee, Laniine, and Gymnorhine. This arrangement was
based not on osteological characters alone, but also on the evidence
of pterylosis and certain wing-muscles. .
The author proposed to include the Vireonide with the Musci-
cape, and the Vireolaniide with the Gymnorhine. With this
last group he proposed, tentatively at any rate, to include the
Paradiseidx, inasmuch as there seemed good reason for continuing
to regard these birds as near allies of the Corvide.
Mr. F. KE. Bepparp, F.R.S., read a paper on the Anatomy of
a Bornean Frog of the genus Megalophrys, with references to
other genera of Batrachia.
Mr. T. A. Cowarp, F.Z.8., communicated a paper on “ The
Winter Habits of the Greater Horseshoe and other Cave-haunting
Bats,” the result of observations made in the Somersetshire
caverns, where at the end of December and beginning of January
he found that the Bats were not in profound sleep, but moved in
the caves and went into the open for food. This food, the author
showed, was not all taken when the Bats were in flight, but was
usually devoured when the Bats were at rest. The manner of
feeding was described and information supplied about the food
of the Greater Horseshoe and the parasites which infested this
species and the Lesser Horseshoe.
The next Meeting of the Society for Scientific Business will be
held on Tuesday, the 23rd April, 1907, at half-past Hight
o'clock P.M., when the following communications will be made :—
1. Mons. 8. A. Bururtry.—On some new or little-known
Siberian Birds.
2. Mr. R. Lypexxer, F.R.S., F.Z.S.—The Ears as a Race-
character in the African Elephant.
3. Mr. Otpriztp Tuomas, F.R.S., F.Z.8.—The Duke of
Bedford’s Zoological Exploration in Eastern Asia.—IV. List of
small Mammals from the Islands of Saghalien and Hokkaido.
With an Appendix on the Cold-blooded Vertebrates by Mr. G.
A. Boutrnesr, F.R.S., F.Z.8.
4. Mr. Henry Scuerrey, F.Z.S.—Some Notes on Hybrid
Bears,
21
The following Papers have been received :—
1. Mr. C. J. Wira.—An Account of the South American
Cheliferine in the Collections of the British and Copenhagen
Museums.
2. Mrs. O. A. Merritt Hawxkes.—On the Abdominal Viscera
and a Vestigial Seventh Branchial Arch in Chlamydoselauchus.
3. Mr. Frank E. Bepparp, F.R.S.—On some new Species of
Eudrilide belonging to the Genera Polytoreutus, Newmaniella,
and Hminoscolex from Mt. Ruwenzori.
4. Mr. Ouprre:D Tuomas, F.R.S.—The Duke of Bedford’s
Exploration in Kastern Asia.—V. Second List of Mammals from
Korea.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL SOCIETY OF LonpDon should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 Hanover Square, Lonpon, W.
April 16th, 1907.
rim ree.
bef iv Ph “a
cht
No. 44.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON,*
April 23rd, 1907.
Dr. J. Rost Braprorp, F.R.S., Vice-President, in the Chair.
-The Srcretary read a report on the additions that had been
made to the Menagerie in March 1907.
Dr. A. Smita Woopwarp, F.R.S., exhibited an antler of a
Red Deer which had become malformed and enlarged by disease.
The specimen was obtained by Mr. Thomas Sheppard from a
prehistoric peat-deposit at Mablethorpe, Lincolnshire.
Mr. R. I. Pocock exhibited, on behalf of the Srcrerary, a model
of the African Elephant “ Jumbo,” formerly living in the Society’s
Menagerie, made by the late Mr. William Prehn and presented to
the Society by his widow.
Mr. R. Lypexxer, F.R.S., read a paper on the ears of the
African Hlephant as a race character, to illustrate which a large
number of photographs and several specimens were exhibited.
The author considered that there must be many more local races
than those already named by Dr. Matschie, although, with the
present material, he hesitated to give separate designations to
several of these. He ventured, however, to propose new names
fer the Elephant of the eastern side of Cape Colony ; for that of
Mashonaland, as typified by a head in the Imperial Institute; for
that of the Lake Rudolf district, as represented by a head pre-
sented to the British Museum by Mr. H. 8. H. Cavendish; and
for the Somali Elephant, as typified by a head in the collection
of S.A.R. le Due d’Orléans at Wood Norton, this last race
being characterised by the very small ears, which, however, were
quite different in shape from those of #. a. knochenhaueri.
* This Abstract is published by the Society at 38 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subseribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Strpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance
24
The author also directed special attention to a skull from the
Albert Nyanza district, for which he had previously suggested
the name /. a. albertensis. Differing in many points from those
of other African elephants, this skull showed a remarkable resem-
blance to that of the extinct Indian Z. planifrons, thus suggesting
the descent of the African Elephant from that species.
Mr. H. E. Dresser, F.Z.S., communicated a paper by Mons. S.
A. ButTuRLIN containing descriptions of three new species and five
new subspecies of Siberian Birds.
Mr. Ouprietp THomas, F.R.S., read a list of small mammals
which had been obtained in the Islands of Saghalien and
Hokkaido by Mr. M. P. Anderson, for the Duke of Bedford’s
exploration of Hastern Asia. Fourteen species were recorded
from Saghalien and thirteen from Hokkaido. The faunas of the
two islands proved to be very similar to each other, although in
some cases subspecific differences between the representative
forms in each were perceptible. In one genus only, Wicromys,
the relationship of Hokkaido seemed to be with the main island
of Japan rather than with Saghalien.
A list of the cold-blooded Vertebrates of Saghalien, by Mr. G.
A. Boutencsr, F.R.S., was also read.
My. Henry Scuerren, F.Z.8., read some notes on Hybrid
Bears, referrmg to cases that had occurred in the Society’s
Gardens, the long series bred by Herr Nill in his Zoological
Garden at Stuttgart (now broken up), and a recent case in
the Garden at Halle-an-der-Saale. Reference was also made
to cases said to have occurred at Cologne and Hanover, but for
these the evidence was not conclusive.
Mr. F. E, Bepparp, F.R.S., contributed a paper on some new
species of Karthworms of the family Eudrilide, belonging to
the genera Polytoreutus, Neumaniella, and Eminoscolex from
Mt. Ruwenzori.
A communication from Mr. C. J. Wrrx contained an account
of the South-American Pseudo-scorpions of the family Cheliferidee
in the collections of the British and Copenhagen Museums.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 7th May, 1907, at half-past Hight
o'clock P.m., when the following communications will be made :—
1. Mr. Ouprizrtp Tuomas, F.R.S.—The Duke of Bedford’s
Exploration in Hastern Asia—V. Second List of Mammals from
Korea,
25
2. Mr. Ausyn R. B. Trevor-Batryr, F.Z.S.—On some Con-
structional Features in Continental Zoological Gardens.
3. Dr. Witt1am E. Hoyviz.—The Marine Fauna of Zanzibar
and East Africa, from Collections made by Cyril Crossland in
1901-1902.—Cephalopoda.
The following Papers have been received :—
1. Mrs. O. A. Murrirr Hawkes.—On the Abdominal Viscera
and a Vestigial Seventh Branchial Arch in Chlamydoselachus.
2. Mr. JAmes Rircurz, M.A.—Some Collections of the Cape
Verde Islands Marine Fauna, made by Cyril Crossland, July to
September 1904.—Hydroids.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL SOCIETY OF LONDON should be addressed to
P. CHALMERS MITCHELL, Secretary.
3 HANovER Square, Lonpon, W.
April 30th, 1907.
‘ y
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ANREP REMCN n
CONTENTS (continued).
‘ April 9, 1907.
Page
Mr. R. I. Pocock. Exhibition of a photograph and the skull of a specimen of Pallas’s
Cat that had recently died in the Menagerie, with general remarks on the species .... 299
I. On a small Collection of Fishes made in the Eastern Watershed of the Transvaal by
Capt. G. E. Bruce. By G. A. Boutunomr, F.R.S.,F.Z.8. (Plates XVIII. & XIX.).. 307
os On the Winter Habits of the Greater Horseshoe, Rhinolophus ferrum-equinum (Schreber),
and other Cave-haunting Bats. By T. A. Cowarp, F.Z.8. 2.0.0.0... ee cece ee ees 312
$
- Notes upon the Anatomy of a Species of Frog of the Genus Megalophrys, with references
to other Genera of Batrachia. By Frank E. Bepparp, M.A., F.R.S., Prosector to
ShiSaSORIOb Yves ssc) kcle oe « sve longs ctu taco Minus teceate BHP Ie ar Peer be Ss teen roger
4. Contributions to the Osteology of Birds.—Part IX. Tyranni; Hirundines; Muscicape,
Lanii, and Gymnorhines. By W.P. Pycnart, F.Z.S., A.L.S., Ge, «22. ce ee esos ee 352
April 23, 1907.
The Secretary. Report on the Additions to the Society’s Menagerie during the month of ©
MarcliwhOO fa emisec nee. s Riek Lun iahe uae RSS ot aaten peace ele ironic pear oD et ae Honma
Dr. A. Smith Woodward, F.R.S., F.Z.S. Exhibition of a malformed antler of the Red Deer. 380
Mr, R. I. Pocock. Exhibition of a model of the African Elephant “Jumbo” .......... 380
Mr. C. J. With. Notice of a Memoir entitled ‘An Account of the South-American
Chelifering in the Collections of the British and Copenhagen Museums ............ 380
J. The Ears as a Race-Character in the African Elephant. By R. LypEKKER .......... 380
2, The Duke of Bedford’s Zoological Exploration in Eastern Asia.—IV. List of small
Mammals from the Islands of Saghalien and Hokkaido. By Oxprimtp Tuomas,
FE.R.S., F.Z.8. (With Appendix on the Cold-blooded Vertebrates, by G. A. BouLencer, ,
TSS EN gS) OR ee eee De SOG ale otra Se eee ontario) 4) «ele, hc) ss 404
#3 0n some new Species of Earthworms of the Family Eudrilide, belonging to the Genera
' Polytoreutus, Newmanniella, and Eminoscolex, from Mt. Ruwenzori. By Frank E.
DEST ARID) I MESAU SHV IR: Surle iuar fo) ee oc ae cay ie enana anal Mare undies DE ar oti hora Died ote heGat ten ats 415
4, Some Notes on Hybrid Bears. By Heyry Scuerren, TZ Sul ys a seer Nate ence ntei atcha cieh tess: 431
BRiried escatctey tape Maree Uae peepee als fo. vals, ¢ lakes veh cu tatelh aria c/a eb eee Mayenne es agai clad euel mie tai ota allot 437
PRUE Gr iarctetercia sist tetaleiarc ose >, oie + o/a:0.8) ocaoieRA eRe ait oye. siex Siintgut aeons ee taeye LalaiLsia0 (ci oye a diet Uae i
List of Council and Officers TC Omi ce APTS sUeare ten Mea ean Tine Aare) > Ce Ca il
List of Contents ....... EAM RE AMA BERTO Obs) Obi TO AR SCO NEY GOO BUS. ORS Wetbeume nna s cS ili
Hiphnabeticalslishot Contributors sess. vesaniay Sees ees eRe comer oni aise ix
Masher Plates * fete oe iia ss. ‘scoala he a eee np Site Anan ena er ce xvii
List of Text-figures .......... eiahakd safuh Os tar taneysl Cage Keir rar nineeren amemua ater! syecata! Sa ahate GP avelern IK
List of New Generic Terns ........ PRA HEe os ta YR RNS Sap ysl ORE A AY NCCT IC AY RS o-ohie
LIST OF “PLATES,
1907, pp. 237-446.
Plate © Page
Deer Hlephant Remains from Crete... ccc cee ee oe we 238
XIV. \ Tanganyika: POly Zone iets «sts ees eiciee's act minielescheuaiciiats 250
Xv. J
XVI. 1. Potamon (Potamonautes) orbitospinus. 2-7. i
Region of various Potamonide .........seeseee eres |
258
XVII. 1&3. Potamon (Potamonautes) platynotus. 2 & 4, Platy- |
thelphusa conculcata.: 5& 6. Platythelphusa maculata .
OVD BaP DUS OF 21620. ie PREBSECLOM wile woo ge ce eickve sue wee eialetae nes
XT Regi conuels. OTUCII Senate es os ioiccs alehec ae Ae wale cere aie aioe J
NOTICE.
The ‘ Proceedings’ for the year are issued in fowr parts, forming two volumes;
as follows:— :
Papers read in January and Fsbruary, in June.
» a Mareh and April, in August.
ss » May and June, in October.
i » November and December, in April,
‘ Proceedings,’ 1907, pp. 1-236, were published on June 12th, 1907.
The Abstracts of the papers read at the Scientific Meetings in
March and April are contained in er Part.
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